THE 


Library  of  tiie  flew  York 
State  Medical  Association. 


LAWS  AID  MECHANICS  OF  CIRCULATION, 


WITH    THE 


PRINCIPLE  INVOLVED  IN  ANIMAL  MOVEMENT. 


WM.   H.  TRIPLETT,    M.  E>., 
Member  of  the  American  Association  for  the  Advancement  of  Science,  etc.,  etc. 


Was  nicht  bewiesen  ist 
Das  braucht  Man  nicht  zu  glauben. 


NEW    YORK  : 

J.   H.   Vail  &   Co., 
1885. 


COPYRIGHTED  BT 

W.    H      TRIPLETT,    M.  D, 
1885. 


ATKIN  &  PROUT,  Printers, 
Nos.  16  and  18  Chambers  Street,  New  York. 


DEDICATION. 


TO 
DR.     ALBERT     GUNTHER, 

The  distinguished  author  and  naturalist,  British  Museum,  in  recognition  of  his 

great  services  as  an  original  investigator,  and  for  his  kindly  heart,  quick 

perception,  and  swift  foot  to  lighten  the  labors  of  others,  which 

rounds  out  and  perfects  character — a  star  in  the  galaxy — 

this  work  is  reverently  and  affectionately  dedicated  by 

The  Author. 


VI  INTKODUCTION. 

anatomy,  very  creditable  to  the  anatomists  and  histologists,  but  of  no  earthly 
use  in  the  p  .ysiologies,  not  being  explained  or  elucidated  as  means  to  ends, 
which  anatomy  purely  is,  cocsisting  of  relative  adjustments  for  special  work, 
since  it  all  relates  to  work,  for  which  they  are  the  relative  adaptations. 

3.  The  Harveian  theory  of  the  circulation,  1628,  but  no  explanation  of  the 
portal  circulation  and  for  maintaining  it  in  correspondence  with  the  absorp- 
tive processes  for  producing  a  balance,  otherwise  impossible ;  no  explanation 
of  the  tissue-circulation  or  that  connected  with  the  cell-brood,  the  objective 
point  for  all  the  commerce,  the  cells  being  the  workmen  in  the  tissues ; 
a  vaso-motor  system  of  nerves,  but  not  competent  to  explain  the  phenomena 
manifested  through  them  ;  no  attempt  to  explain  the  numerous  muscles  and 
nerves  (unknown  to  Harvey)  to  the  vessels. 

4.  Numerous  physiological  experiments  upon  the  nerves,  muscles,  blood- 
vessels, etc.,  by  means  of  which  deeply  interesting  phenomena  are  developed, 
but  their  relevancy  to  any  fundamental  circumstance  in  the  organism,  or  con- 
nection with  one  another  by  reason  of  any  principle  in  the  mechanics,  not 
shown  ;  illustrated  by  explanatory  cuts  showing  the  mode  of  procedure. 

Traube's  Curves  are  beyond  its  reach  ;  so  also  the  blood-pressure  curves 
and  the  curves  of  intra-thora  pressure.  It  offers  no  solution  for  the  existence 
of  "  inhibitor"  and  "  accelerator"  nerves  to  the  heart.  It  cannot  explain  the 
anatomy  in  the  intestines,  and  why  there  should  be  a  diaphragm  in  the  mam- 
malia only.  It  cannot  get  fat,  albumen,  alcohol  or  any  non-dializable  sub- 
stance into  the  body,  yet  they  are  rapidly  absorbed  and  the  explanation  easy. 

More  or  less  remarks  upon  muscular  irritability  and  contractility,  but  noth- 
ing of  dualism  in  the  muscles,  without  which  musculation  would  be  utterly 
impossible,  etc.,  etc.  The  books  large  enough,  too  ;  some  of  them  a  thousand 
pages,  nearly,  fine  press.  No  grasp,  no  continuity  of  relations,  all  differing, 
but  as  far  as  possible  from  a  philosophy.  Each  one  following  the  train  of  facts 
he  is  most  familiar  with  (compelled  to  it  in  the  absence  of  a  guiding  principle), 
fitting  the  others  into  the  text  as  best  he  can  to  make  a  book,  at  times  appar- 
ently as  though  tumbled  in  at  random  till  the  book  is  big  enough;  vain 
strivings  for  order  and  method,  however,  discernible.  The  result  nil.  (I  would 
like  to  make  exceptions,  but  «annot ;  though  some  of  the  books  have  remarka- 
ble excellence,  and  it  wounds  one  to  the  heart  to  say  aught  against  them.) 
With  all  this,  wrong  teaching  most  pernicious,  doing  irreparable  harm  to 
some  minds,  worse  than  night-blindness,  for  that  would  suffer  the  unfortunate 
to  see  by  dayMght,  but  this  by  no  light  whatever ;  but  to  any  mind  the  work  of 
wnlearning  and  starting  afresh  is  most  difficult,  next  to  impossible,  accomplished 
only  by  persistent  and  heroic  effort.  For  example,  respiration  and  circulation 
form  a  connected  movement  for  supplying  the  cell- brood  in  the  tissues  and 
removing  waste  products  ;  but  the  physiologies  treat  them  separately,  basing 
circulation  upon  the  heart,  respiration  upon  the  diaphragm.  By  what  right  is 
that  done  ?    No  right ;  none. 

The  birds,  reptilia,  and  the  rest  of  them,  have  no  diaphragm  ;  neverthe- 
less, they  respire  ;  besides,  produce  vocal  resonance  characteristic  of  them, 
showing  that  respiration  is  also  under  voluntary  control,  the  same  as  in  mam- 
malia. Air  is  got  into  the  lungs  by  means  of  a  pleural  vacuum — speaking 
more  correctly,  a  "  negative  pressure  in  the  pleurae" — distending  the  organs 
in  the  first  place  by  this  means,  then  producing  inspiration  by  contracting  the 


INTRODUCTION.  Vll 

diaphragm  and  intercostal  muscles,  then  expiration  by  relaxing  the  muscles. 
It  will  not  answer,  and  is  not  the  method,  as  is  fully  proven  in  the  text  by  the 
most  incontrovertible  evidence.  Furthermore,  the  birds,  etc.,  have  no  such 
explanation,  passing  them  by  ;  but  this  cannot  be  done  in  physiology  ;  it  all 
must  be  explained.  And  in  the  case  of  the  worms,  we  have  a  complex  circu- 
lation carried  on  where  there  is  no  heart,  the  vessels  at  the  same  time  expand- 
ing and  contracting  regularly  and  rhythmically,  possessing  automatism  the 
same  as  the  heart,  being  also  furnished  with  muscles  and  nerves  for  increasing 
force  in  them  commensurate  with  the  physiological  requirements,  You  can- 
not begin  at  the  end  of  animal  life  to  write  about  circulation  any  more  than 
respiration.  In  the  embryo,  but  for  the  power  to  produce  circulation  inde- 
pendently of  the  heart,  the  heart  itself  could  not  be  developed,  while  vascular 
development  goes  on  simultaneously,  the  intestinal  canal  and  other  organs 
also,  the  heart-force  coming  in  subsequently  for  increasing  the  action  simply, 
the  vessels  growing  pari  passu  with  it.  The  theory  breaks  down  and  will  not 
answer  the  physiological  requirements  ;  is  therefore  incorrect.  Then,  again, 
the  heart  is  used  as  a  force-pump,  which  involves  a  suction  with  a  driving 
force,  but  entirely  ignoring  the  former  by  using  contraction  in  the  cardiac 
muscles  simply  ;  whereas  the  organ  would  first  have  to  be  filled  by  diastole 
before  one  may  speak  of  forcing  the  blood  out  of  it  by  systole,  thus  producing 
a  current  through  the  organ,  while  expansion  and  contraction  are  correlated 
forces  in  Nature,  therefore  inseparable. 

It  is  needless  to  extend  the  matter,  for  physiology  could  not  be  more  impotent ; 
nor  can  help  come  by  scalpel  and  microscope  :  their  work  has  been  well  done, 
the  body  taken  down  to  the  very  cells  themselves  and  polarizing  lenses  are  set 
to  them.  In  short,  analysis  may  be  said  to  be  complete,  but  the  key  to  animal 
structure  and  function  not  forthcoming,  not  to  be  forced  by  scalpel  or 
microscope,  or  rather  the  latter,  for  the  work  is  now  in  micro-organisms  and 
cells.  Nor  can  physiology  remain  where  it  is.  The  ground  sinks,  no  firm 
foothold,  every  effort  but  sinking  you  deeper  in  the  morass.  A  change  must 
come,  else  there  is  no  hope.  The  key  to  be  found  at  all?  Of  course  it 
should.  The  arrangements  are  visible  and  persistent,  and  for  their  full  and 
complete  interpretation  requires  only  that  the  proper  method  be  adopted. 

Set  physiology  in  the  opposite  direction,  or  toward  synthesis  and  reconstruc- 
tion, and  you  ivill  find  it,  for  this  will  take  you  to  the  organic  law  upon  which 
the  animal  organism  is  based,  and  that  is  the  key  ;  for  it  is  as  utterly  impossi- 
ble to  explain  the  mechanics  in  the  absence  of  this  principle  as  to  explain  any 
mechanism  in  the  absence  of  the  principle  it  involves,  and  the  animal  body  is 
a  mechanism.  And  though  it  differs  from  all  others  in  that  it  is  automatic, 
nevertheless  this  would  not  do  away  with  the  necessity  for  a  principle  upon 
which  to  base  the  mechanics,  as  must  appear  obvious. 

Life  so  mysrerious  !  Granted.  But  life  works  in  matter,  and  matter  has 
laws  eternal  and  inflexible  regulating  it.  What  law  have  you,  then,  as  the 
basis  of  animal  mechanics  with  which  everything  in  the  organism  must  have 
adjustment  and  every  piece  its  place  and  relative  value,  in  the  very  nature  of 
things  ?  None.  Then,  it  is  manifest  that  you  cannot  explain  the  mechanism  or 
write  a  physiology.  In  order  to  do  this,  the  principle  would  have  to  be  forth- 
coming first,  and  from  that  stand-point  it  is  all  made  intelligible  ;  otherwise,  is 
utterly  inexplicable.     But  you  cannot  begin  with  animal  or  even  floral  life,  for 


Vlll  INTRODUCTION. 

the  principle  is  not  confined  or  pent  up  in  them,  extending  into  the  very  me- 
chanics for  producing  circulation  in  air  and  water,  out  of  which  living  organ- 
isms are  evolved.  Commencing  with  this,  thence  through  flora  and  fauna, 
noting  the  method  for  increasing  it  in  the  latter  commensurate  with  the  force 
expended  in  them  ;  when  it  will  turn  out  that  physiology  is  but  natural  philos- 
ophy, aDatomy  means  to  ends,  the  functions  resultant  phenomena ;  or,  in 
other  words,  physiology  is  the  descriptive  anatomy  and  functions  in  the  organs, 
the  former  being  the  special  adaptations  of  means  to  ends  in  correspondence 
with  the  character  and  amount  of  work  performed,  since  it  all  relates  to  work 
which  must  have  its  equivalent  in  force  there  and  then ;  the  whole  evolved 
out  of  circulation,  while  this  in  turn  is  regulated  by  law  which  is  being  inces- 
santly invoked  in  the  measure  of  the  requirements.  Comprehensive  it  is,  but 
easily  understood. 

Finally,  why  should  it  be  deemed  an  incredible  circumstance  to  have  a  fun- 
damental principle  in  animal  life,  upon  which  the  phenomena  are  based, 
seeing  that  everything  in  the  visible  universe  is  regulated  by  law,  and  that 
there  is  systematic  and  persistent  arrangements  in  the  structures  ;  otherwise 
inexplicable?  Furthermore,  method  implies  principle.  And  when  similar 
arrangements  in  the  structures  and  movements  in  the  parts  habitually  occur 
in  great  classes  of  animals,  we  may  rest  assured  that  a  common  principle 
underlies  it  all ;  and  in  order  to  interpret  the  phenomena  it  would  be  abso- 
lutely necessary  to  first  make  out  the  mechanical  principle  that  applies.  And 
since  the  whole  relates  to  work,  there  should  be  no  difficulty  in  determining 
this  circumstance. 

Such,  in  brief,  is  the  course  adopted  in  this  work,  commencing  with  the  laws 
and  mechanics  of  circulation  as  applied  to  the  pumping  movements  in  lungs, 
heart  and  vessels,  which  undoubtedly  relate  to  rhythmical  changes  in  pressure 
for  increasing  circulation  commensurate  with  the  force  expended  in  the 
organism,  seeing  also  that  these  movements  are  in  correspondence  with  the 
activities,  rising  and  falling  with  the  swell  in  the  activities,  the  whole  forming 
a  connected  movement  in  the  very  nature  of  things.  And  while  there  is 
automatism  in  the  organs  and  the  heart  continues  to  beat  for  a  time  after  res- 
piration is  suspended,  it  is  no  infraction  of  this  principle,  for  in  no  other  way 
could  a  balance  be  maintained  in  the  organism,  the  heart  and  vessels  being 
simply  a  carrier  between  the  cell-brood  and  lungs,  while  respiration  is  the 
great  pumping  action  for  compelling  the  commerce  in  the  organism.  In 
short,  that  respiration  answers  to  the  pendulum  in  the  clockwork,  while  the 
heart  and  vessels  are  set  to  this  as  the  minute  hand  to  the  hour  hand  ;  so  that 
it  all  works  together  harmoniously  and  in  perfect  concord,  to  the  end  that  a 
stream  of  commerce  may  be  maintained  between  the  cell-brood  and  environ- 
ment, from  which  all  the  supplies  are  obtained,  and  into  which  in  due  time 
waste  products  are  returned  for  redistribution,  the  eternal  order  of  tilings. 

Beginning,  then,  with  the  principle  for  producing  circulation  in  air  and 
water,  and  for  pulling  the  molecules  into  position  and  locking  them  in  the 
structures,  or  similar  to  what  takes  place  in  crystal  structure  (which  is  ap- 
proximated in  plants  and  bones,  while  in  the  soft  tissues  there  is  provision  for 
freer  molecular  action)  you  come  into  immediate  contact  with  that  all-absorb- 
ing question,  What  is  Life  ?  I  cannot  make  out  aught  else  than  a  form  of  force 
— wonderful  metamorphoses  in  force  ;    using  present  terminology,  Life  is  an 


INTRODUCTION".  IX 

"eddy" in  force.  With  force,  itself ,  there  we  stop.  The  ultima  Thule !  But 
this  we  know  of  a  surety,  that  force  is  controlled  by  laws  eternal  and  inflex- 
ible, while  matter  manifests  them,  and  is  energized  by  force.  Nor  need  we 
fear  to  follow  to  their  logical  conclusions  the  fundamental  principles  in  Nature. 
A  stream  cannot  rise  higher  than  its  source  ;  and  man  intelligent,  it  follows 
there  must  necessarily  be  intelligence  in  Nature — all -pervading  intelligence  ; 
further  I  cannot  say  ;  das  ist  genug. 

In  conclusion  :  This  work,  as  the  name  implies — Life  :  Its  Laws  and  Mechanics 
— would  include  all  vital  phenomena,  but  as  circulation  underlies  it  all  (fur- 
nishing the  materials  for  elaborating  structure  and  evolving  force,  but  involv- 
ing the  principle  concerned  in  animal  movement,  in  order  to  increase  circula- 
tion), this  would  naturally  have  precedence,  the  rest  following  in  regular 
order,  to  be  published  as  a  serial,  beginning  with  "Gravitation  and  Devel- 
opment," gravitation  being  pivotal,  so  to  speak,  in  development,  showing 
the  intimate  relations  it  sustains  to  the  latter  to  the  very  minutia  in  tissue- 
structure,  the  organs  of  circulation  with  those  of  respiration,  locomotion, 
etc.,  but  most  manifest  in  the  frame- work  of  support  to  the  soft  tissues  or 
bony  skeleton  ;  a  very  notable  circumstance  being  the  mode  of  conserving 
the  central  nervous  system  by  supporting  it  upon  a  ' '  water-bed"  formed  of 
the  cerebro-spinal  fluid  and  the  fibrous  leaflets  in  the  "falx  cerebri" 
and  "  tentorium  cerebelli,"  with  the  nerves  running  into  them  for  regu- 
lating tension,  so  as  to  obviate  concussion  in  locomotion,  leaping,  etc. ; 
the  principle  involving  the  articulations  themselves,  for  the  brain  must  be 
protected  against  jar.  But  all  the  organs  exhibit  adjustments  with  gravi- 
tation. The  great  development  of  the  vascu]ar  system  in  land  animals, 
the  numbers  of  the  muscles  and  nerves  in  the  heait  and  vessels,  have 
similar  explanation,  the  fluids  tending  constantly  to  the  earth.  In  fine, 
every  movement  in  the  mass  and  molecules  must  necessarily  involve  a 
struggle  with  this  force,  as  must  appear  obvious.  Nay,  to  the  very  mode  of 
rising  up  and  lying  down,  the  animals  doing  it  differently  by  reason  of  the 
special  adjustments  that  obtain  in  the  viscera,  the  ruminants  elevating  the 
hinder  par:  s  first,  then  bringing  up  the  fore  parts,  reversing  the  action  when 
lying  down ;  whereas,  the  other  animals  elevate  the  fore  parts  first,  then 
bringing  up  the  hinder  parts,  reversing  the  action  when  lying  down.  And 
that  it  is  not  awkwardness  in  the  ruminant  which  causes  it  to  take  the 
course  it  does,  is  proven  by  the  fact  that  the  fleet  deer  and  nimble  antelope  go 
through  the  same  movements.  Finally,  strange  and  fanciful  as  it  may  seem, 
nevertheless,  the  relative  stages  in  development  in  the  anthropoid  apes  are 
readily  determined  by  it,  beyond  the  shadow  of  a  doubt,  and  gorilla  is  closest 
to  man  of  any  existing  species,  which  is  not  a  figment  of  the  mind  born  of  a 
lively  imagination,  but  a  sober  fact  wrought  in  the  bony  skeletons  of  the  ani- 
mals, very  plain  and  easily  to  be  seen.  We  would  not  anticipate,  however,  but 
let  the  present  work  serve  as  earnest  of  that  engagement. 

Nature  works  by  laws  immutable,  and  everything  is  determined  bylaw,  while 
order  and  method  are  made  inevitable.  And  the  very  fact  itself  that  so  much 
is  explained  within  the  compass  of  the  present  work,  is  conclusive  proof  that 
correct  Lines  have  been  adopted. 


Digitized  by  the  Internet  Archive 

in  2010  with  funding  from 

Open  Knowledge  Commons  (for  the  Medical  Heritage  Library  project) 


http://www.archive.org/details/lawsmechanicsofcOOtrip 


Ubrarj  of  iiko  TSTewTork 

State  Medical  Association. 

CONTENTS. 


CHAPTER    I. 

MECHANICS  FOR  PRODUCING  CIRCULATION  IN  AIR  AND  WATER, 
AND  FOR  CONNECTING  THEM  WITH  CIRCULATION  IN  LIVING 
ORGANISMS. 

The  Fundamental  Fact  at  the  Basis  of  Living  Organisms— Living  Organisms  Evolved 
from  Substances  Contained  in  Air  and  Water— Mode  of  Suspending  Atmospheric 
Matter  in  Space,  and  the  Principle  for  Producing  Movement  among  the  Mole- 
cules—Experiments of  Dufay  and  Faraday  with  Electricity  and  Magnetism— Action 
of  Gravitation  Overcome  by  Electrical  Force— Transformations  of  Heat  into  Elec- 
trical Force  for  Energizing  the  Polar  Forces  among  the  Molecules,  the  Fundamental 
Principle  for  Producing  Circulation  in  Air  and  "Water— The  Prodigious  Force  which 
is  Involved  for  Producing  Circulation  in  the  Atmosphere  and  in  "Water— The 
Mechanics  Connected  Through  and  Through  by  Means  of  Electrical  Force- 
Electrical  Phenomena  in  Storms— Mode  of  Effecting  Changes  in  Pressure  for 
Increasing  Circulation  in  the  Atmosphere— The  Action  in  "Water— Principle  in 
Diffusion,  Osmose,  Capillarity— Organic  Connection  Subsisting  between  Circula- 
tion in  Air  and  Water  and  in  the  Living  Organism        ...       -       Page  1 

CHAPTER    II. 

THE  FLORAL  CIRCULATION. 

Principle  in  the  Floral  Circulation — Action  of  the  Polar  Forces  and  the  Mode  of  Increas- 
ing it — Physiological  Experiment  Demonstrating  the  Energy  of  the  Polar  Forces  in 
Flora — Circulation  in  Flora  Comparatively  Slow — Deduction  to  be  Drawn  There- 
from— Explanation  for  the  Rapidity  of  Circulation  in  Fauna  and  the  Amount  of 
Food  Consumed  by  Them        .----------S3 

CHAPTER    III. 

THE  ANIMAL  CIRCULATION. 

Principle  in  the  Animal  Circulation — Adjustment  with  Pressure  and  the  Power  of  Pro- 
ducing Rapid  Rhythmical  Changes  in  Pressure,  the  Law  in  the  Animal 
Circulation — The  Movements  in  Respiration  :  Heart,  Arteries,  etc.,  Pumping- 
Aetions  for  Increasing  Circulation,  the  Whole  Forming  a  Connected  Movement  for 
Increasing  Circulation  between  the  Cell-Brood  and  Environment,  whence  the  Nutri- 
tive and  Force-Producing  Elements  are  Derived,  and  into  which  the  Waste  Products 


1  CONTENTS. 

are  Returned — Explanation  for  the  Correlation  of  the  Vaso-Motor  and  Voluntary- 
Motor  Centres  with  the  Respiratory  Centre — Mechanics  in  Inspiration  and  Expira- 
tion— Two  Respiratory  Movements  Performing  at  the  Same  Time  ;  One  in  the 
Lungs,  the  Other  in  the  Tissues — Physiological  Experiments,  Showing  that  Pressure 
is  the  Fundamental  Circumstance  in  the  Animal  Organism  with  which  Everything 
has  Adjustment,  and  that  the  Actions  in  the  Lungs%  Heart,  Vessels  and  Hollow 
Viscera  Relate  to  Changes  in  Pressure  for  Compelling  Movement  in  the->Contents  : 
t.  e.,  for  Increasing  Circulation— Otherwise  are  Meaningless  29 


CHAPTER    IV. 

RESPIRATION  IN  DIFFERENT  STAGES  IN  DEVELOPMENT. 

Import  of  Amoebae  Movement — The  Alternate  Extension  and  Retraction  of  the  Branched 
Processes,  a  Pumping  Action  for  Increasing  Circulation — Why  Locomotion  Should 
Increase  Circulation  Correspondingly — The  Pumping  Movements  Analogous  with 
Respiration — The  Action  in  Vacuoles  and  the  Radiating  Canals,  an  Early  Indication 
of  the  Mechanical  Principle  in  the  Heart  ancLArteries,  the  Latter  Expanding  as  the 
Former  is  Contracting  and  Vice  Versa — The  Action  in  Gastrula — Necessity  for  Co- 
ordinating the  Mucous  with  the  Skin-Surface,  in  Order  to  Produce  Afflux  and 
Efflux  of  the  Fluids  in  the  Body-Interior — This  Circumstance  Further  Dlustrated  in 
the  "Worms  for  Producing  the  Undulations  which  Course  along  the  Body  During 
Imbibition— The  Principle  Applied  to  Respiration  and  the  Action  Taking  Place  in 
the  Lungs — Illustrated  in  the  Prog,  in  which  it  is  Demonstrated  Experimentally 
that  the  Lungs  Expand  and  Contract  Regularly  and  Rhythmically  Synchronous 
with  the  Action  Taking  Place  in  the  Muscular  Envelope  or  Containing  Walls,  in 
Order  to  Produce  Afflux  and  Efflux  of  Air  and  Blood  in  these  Organs  for  Respira- 
tory Purposes ;  Otherwise  Impossible — The  Manner  the  Parts  are  Coordinated — 
Dependence  of  the  Portal  Circulation  upon  Respiration — the  Same  Principle  in 
Mechanics  for  Every  Stage  in  Development — The  Action  in  Birds — The  Special 
Adjustments  in  the  Viscera — Portal  Circulation  in  -        4G 

CHAPTER    V. 

RESPIRATION  IN  MAMMALIA. 

Respiration  in  the  Mammalia  Fundamentally  the  Same  as  in  Birds  and  Reptiles,  the  Lungs 
and  the  Muscles  in  the  Abdomen  Expanding  and  Contracting  Simultaneously — Office 
of  the  Diaphragm  and  the  Ocassion  for  it — The  Action  in  the  Diaphragm  Alternating 
with  the  Muscles  in  the  Abdomen,  the  One  Contracting  as  the  Other  is  Expanding,  and 
Vice  Versa — The  Action  the  Diaphragm  Exerts  upon  the  Ribs,  Bending  and  Flaring 
them  Open  upon  the  Sides  During  Inspiration,  and  Vice  Versa  During  Expiration — 
Mode  of  Coordination,  Inclusive  of  the  Lungs,  the  Whole  Moving  in  Perfect  Con- 
cert— Circumstances  in  the  Vital  Phenomena,  Anatonncal  and  Physiological,  which 
Make  it  Absolutely  Certain  that  the  Lungs  Expand  and  Contract  Regularly  and 
Rhythmically  Synchronous  with  the  Actions  in  the  Diaphragm  and  Containing 
Walls— Physiological  Experiments  Proving  this  Circumstance — The  Action  in  the 
Tracheal  System — Mode  of  Maintaining  Cleanliness  in  the  Alveoli  and  Air-Pas- 
sages— Significance  of  a  Cough — The  Action  of  the  Trachea  in  Vocalization — Ex- 
planation for  the  Devious  Course  of  the  Recurrent  Laryngeal  Nerves,  which  First 
Descend  into  the  Chest  to  Connect  with  the  Lungs  Before  Proceeding  to  the  Larynx 
and  Vocal  Cords,  A  scending  thence  Upon  the  Trachea  for  this  Purpose,  no  matter 
how  Long  the  Neck  may  be  :  e.  g.,  the  Giraffe 83 


CONTENTS.  Xlll 

CHAPTER    VI. 

THE  MECHANICS  FOR  CIRCULATING  BLOOD  IN  THE  LUNGS,  AND  THE 
ACTION  OF  THE  HEART  AND  ARTERIAL  SYSTEM  IN  CONNECTION 
THEREWITH. 

Mechanics  for  Circulating  Blood  in  the  Alveoli — Anatomical  Dispositions  in  the  Walls  of 
the  Alveoli  for  Effecting  it,  or  for  Compelling  Circulation  of  Blood  in  the  Plexuses 
to  be  in  Correspondence  with  the  Circulation  of  Air  in  the  Alveoli — Extent  of  the- 
Alveolar  Plexuses  and  the  Manner  they  are  Affected  by  Inspiration  and  Expiration 
— Functions  of  Residual-Air  in  this  Connection — An  Elastic  Cushion  for  Transmit- 
ting the  For^e  in  the  Lungs  and  the  Muscular  Envelope  upon  the  Plexuses  for  Com- 
pelling the  Blood  to  be  in  Correspondence  with  the  Circulation  of  Air  in  the  Alveolar- 
Chambers — Manner  of  Maintaining  a  Balance  in  the  Dual  Circulations  in  the  Alveoli 
— Relations  which  the  Heart  Sustains  to  the  Pulmonic  Circulation — Mechanical  Prin- 
ciple in  the  Heart  Itself — Anatomical  Dispositions  in  the  Right  Side  Adjustments. 
with  the  Functions  in  the  Lungs— Ditto,  Left  Side — Nerves  for  Effecting  Coordina- 
tion with  the  Lungs — Action  in  the  Arterial  System  Synchronous  with  Respiration 
—Nerves  for  Effecting  it — Physiological  Problem  Connected  with  the  Curves  in 
Blood-Pressure  Tracings  and  the  Curves  in  Intra-thoracic  Pressure— Traube's 
Curves — Physiological  Experiments  Proving  the  Existence  of  Rhythmical  Expan- 
sions and  Contractions  in  the  Arterial  System  Synchronous  with  the  Actions  in  the 
Heart  and  Lungs    -------------        ]  21 

CHAPTER    VII. 

AUTOMATISM  IN  THE  VESSELS,   AS    WELL  AS   IN  THE  HEART. 

Principle  in  Cardo-Arterial  Movement — The  Arterial  System  Expands  as  the  Heart 
Contracts,  and  Vice  Versa — Mode  of  Proving  this  Action  in  the  Vessels — Evidence 
in  Arterial  Tracings — Arterial  Tracings  Contrasted  with  the  Artificial  Tracings, 
Produced  by  the  Apparatus  of  Marey,  Showing  Essential  Points  of  Difference,  and 
Proving  Incontrovertibly  Automatism  in  the  Vessels— The  Variations  in  Blood- 
Pressure  Tracings  Corroborative  of  this  Action  in  the  Vessels,  Confirming  the  De- 
ductions from  Arterial  Tracings — All  the  Elements  in  Arterial  Tracings  and  the 
Tracings  which  are  produced  by  Variations  in  Blood-Pressure  Harmonize — The 
Facts  in  Development  Confirmatory  of  Automatism  in  the  Vessels — Regular  Rhyth- 
mical Expansions  and  Contractions  Taking  Place  in  the  Vessels  of  the  "Worms  in  the 
Entire  Absence  of  a  Heart  for  Producing  Them,  the  Vessels  Expanding  and  Con- 
tracting Themselves— The  Pulsations  in  the  Umbilical  Cord  not  Synchronous  with 
the  Action  in  the  Fcetal  or  Maternal  Heart  ;  Moreover,  Pulsates  even  after  Connec- 
tion is  severed — The  Pulsations  in  the  Ears  of  a  Rabbit  and  in  the  Wings  of  a  Bat 
not  Synchronous  with  the  Action  in  the  Heart,  Showing  Independent  Action  in  the 
Vessels — The  Special  Anatomy  in  the  Vessels  Fundamentally  the  Same  as  in  the 
Heart — Progressive  Increase  of  the  Muscles  in  the  Arterial  System  from  the  Heart 
to  the  Tissue-Territories — Function  of  the  Strong  Elastic  Coat — How  High  Pressure 
is  Produced,  and  the  Occasion  for  it — How  Reflux  in  the  Capillaries  is  Obviated — 
Mode  of  Increasing  and  Diminishing  the  Local  Actions,  and  for  Coordinating  the 
Vessels  with  Respiration  and  the  Action  in  the  Heart.       -----        138. 

CHAPTER    VIII. 

RESPIRATION  IN  THE  TISSUES  AND    THE  ACTION   OF    THE   CAPILLA- 
RIES IN  CONNECTION  THEREWITH. 

Two  Respiratory  Movements  Going  on  at  the  Same  Time  in  the  Body,  One  in  the  Lungs, 


XIV  CONTENTS. 

the  Other  in  the  Tissues — The  Force  for  Producing  the  Movements  Propagated  from 
the  Medulla  Oblongata  and  Respiratory  Centre — The  Composite  Character  in  Arterial 
Tracings  Readily  Explained — Rhythmical  Expansions  and  Contractions  in  the  Tissues 
Synchronous  with  the  Actions  in  the  Lungs,  Heart,  Arteries  and  Capillaries — The 
Waves  Superposed  upon  One  Another  in  the  Order  Named,  or  the  Cardo- Arterial 
upon  the  Respiratory,  and  the  Capillary  or  Dicrotic  Waves  upon  the  Cardo- 
Arterial — Capillary  Action  the  Source  of  Dicrotism — Mode  of  Demonstrating 
this  Circumstance — The  Action  in  the  Capillaries  Producing  a  Current  Into  and 
Out  of  the  Tissue-Interstices — Relations  of  the  Cell-Brood  to  this  Circulation — 
Nervous  Apparatus  for  Connecting  Them  with  the  Capillaries  and  Central  Nerv- 
ous System  for  Increasing  and  Diminishing  the  Local  Circulation,  with  the 
Exigencies  in  the  Functions — Mechanics  in  Blushing — Explanation  for  Arrest  of 
Arterial  Pulsations  upon  the  Distal  Side  in  Aneurisms — Action  in  the  Venous 
System — Functions  of  the  Muscles  and  Nerves  in  Veins — The  Relations  they  Sus- 
tain to  Respiration — Explanation  for  the  Great  Volume  of  Venous  Blood  and  the 
Slowness  in  this  Circulation — Also  for  the  Motion  in  the  Brain  Synchronous  with 
Respiration.     ------ 159 

CHAPTER    IX. 

THE  PORTAL   CIRCULATION  AND    THE  MECHANICS  IN  THE  ABDOMEN. 

The  Intestines  of  Mammalia  Filled  and  Distended  with  Air  for  Increasing  the  Digestive 
and  Absorptive  Processes,  Serving  the  Purpose  of  an  Elastic  Cushion  for  Trans- 
mitting the  Force  in  the  Gut  upon  the  Food  for  the  Purpose — The  Action  not  Un- 
like that  which  Obtains  in  a  Churn — Necessity  for  a  Diaphragm — Existence  of  High 
.  Pressure  in  the  General  Cavity  of  the  Abdomen  Produced  by  the  Air  in  the  Intes- 
tines for  Increasing  the  Portal  Circulation  so  as  to  Maintain  this  in  Correspondence 
with  the  Absorptive  Processes — Principle  of  Coordination  Applied  to  the  Stomach 
and  Walls  of  the  Abdomen  for  Effecting  Ingestion  and  Office  of  the  Pneumogastric 
and  Phrenic  Nerves  in  Connection  Therewith — Why  the  Animal  Rises  when  Eating 
and  Drinking — Why  Respiration  is  Suspended  During  Deglutition  or  After  the  Ali- 
ment has  Passed  the  Glottis,  and  the  Danger  of  Intrusion  in  the  Air-Passages  is  Over 
— The  Action  in  the  Stomach  and  the  Physiological  Anatomy  in  the  Organ — The 
Role  in  the  Air-Cushion  in  Connection  Therewith — Similar  Survey  of  the  Intestines — 
The  Rapid  Absorption  of  Fat,  Alcohol  and  Other  Non-Dializable  Substances  Easily 
Explained — The  Fine  Adjustments  in  the  Muscularis  Mucosae  and  the  Mode  of 
Coordinating  Them  with  the  Muscles  in  the  Walls  for  Making  Absorption  Very 
Effective — No  Difficulty  in  Absorption 185 

CHAPTER    X  . 

RESPIRATION  AND   THE  PORTAL  CIRCULATION. 

Circulation  in  the  Liver  Dependent  upon  Respiration— Mechanics  in  the  Diaphragm  and 
Walls  of  the  Abdomen  Respecting  it — The  Mesentery  a  Soft,  Elastic  Cushion  for 
Effecting  Gentle  Compression  of  the  Liver-Substance,  under  the  Action  in  the  Dia- 
phragm and  Walls  of  the  Abdomen  during  Inspiration  for  Increasing  its  Circulation 
— Mode  of  Demonstrating  this  Circumstance — Effect  upon  the  Portal  Vessels  and 
Lower  Cava-System — Absence  of  Valves  in  the  Veins  within  the  Abdomen,  save  in 
the  Pelvic  Viscera  Only — Explanation  for  the  Latter  Circumstance — Physiological 
Anatomy  of  the  Liver  with  Reference  to  Circulation— Why  the  Hepatic  Veins  are 
Incorporated  with  the  Liver-Substance — Relations  of  the  Portal  System  to  the 
Hepatic  Veins— Why  Rhythmical  Compression  of  the  Liver-Substance  during  Res- 


CONTENTS.  XV 

piration  should  Increase  Circulation  in  the  Inter-  and  Intra-Lobular  Vessels  Oorre- 
pondingly — Automatic  Action  in  the  Portal  Vessels  Synchronous  with  Respiration 
for  Further  Increasing  it — Mechanics  for  Circulating  Bile — The  Action  in  the  Gall- 
Bladder  and  Bile-Ducts — The  Action  in  the  Duodenum,  in  Connection  with  the 
Biliary  and  Pancreatic  Secretions — Adjustments  in  the  Viscera  Necessitated  by  the 
Action  in  the  Diaphragm — Mechanics  Connected  with  the  Openings  in  the  Dia- 
phragm— (Esophagus  Constricted  during  Inspiration,  while  the  Vena-Cava  Lumen 
is  thrown  Widely  Open;  as  also  the  Aortic — Elongation  and  Contraction  of 
(Esophagus  with  Inspiration  and  Expiration — Ditto  Venae  Cava? — Ditto  Portal 
Vessels  and  Renal  Veins — Similarity  in  the  Anatomical  Dispositions  of  the  Muscles 
in  these  Organs      -------___.__         ggQ 


CHAPTER    XI. 

AUTOMATISM    IN     THE     LYMPHATICS     AND     THE     MECHANICS     FOR 
CIRCULATING  LYMPH 

The  Lymphatic  System  Intercalated  between  the  Arterial  and  Venous  Systems, 
Arising  in  the  Tissues  and  Debouching  in  the  Veins  at  the  Root  of  the  Neck — 
The  Dual  Functions  in  these  Organs  Concerned  in  Drainage  and  Haamatosis — 
The  Vessels  Connecting  with  Respiratory  Movement  in  the  Tissues,  Expanding 
and  Contracting  with  the  Blood-Capillaries  by  Means  of  the  Nervous  Connections 
Subsisting  between  Them — The  Action  in  the  Larger  Vessels  and  Gland-Structures — 
Muscles  and  Nerves  to  the  Organs  for  Producing  the  Actions — The  Action  in  the 
Lacteal  System  and  the  Manner  in  which  it  is  Affected  by  the  Action  in  the  Gut 
for  Compelling  Rapid  Movement  of  the  Fluids  in  the  Vessels— The  Relations  which 
this  Sustains  to  Inspiration  and  the  Action  in  the  Venous  System — Physiological 
Experiments  Proving  that  Inspiration  Pumps  the  Lymph  into  the  Venous  System 
Simultaneously  with  the  Portal  and  Hepatic  Blood— Proof  of  Automatism  in  the 
Vessels  -----____._  ..        347 

CHAPTER    XII. 

NERVES  TO  THE  VISCERA  IN  THE  ABDOMEN,  AND  MODE  OF  CONNECT- 
ING THEM  WITH  THE  CEREBROSPINAL  AXIS. 

The  Double  Ganglionic  Dorsal  Chain  in  Vertebrates,  the  Analogue  of  the  Double 
Ganglionic  Chain  in  the  "Worms— Nerves  of  Meissner— Nerves  of  Auerbach— Mode 
of  Connecting  them  with  the  Solar  Plexus  and  Central  Nervous  System— Relations 
of  the  Nerves  of  Meissner  and  Auerbach  with  the  Intestinal  Mucous  Membrane  and 
Epithelium— Mode  of  Controlling  the  Blood-Supply  from  the  Aorta-Trunk  by  Means 
of  the  Cceliac  Axis,  Superior  and  Inferior  Mesenteric  Arteries  and  their  Branches,  so 
that  Every  Organ  and  Fractional  Portion  of  the  Same  can  Regulate  their  own  Sup- 
plies in  the  Measure  of  the  Functional  Activities— Connection  of  the  Pneumogastric 
Nerves  with  the  Solar  Plexus— Mode  of  Connecting  the  Solar  Plexus  and  Spinal 
Ganglia  with  the  Dorsal  Nerves  and  Spinal  Medulla,  the  Nerves  of  the  Ganglionic 
Chain  running  up  Both  Roots  of  the  Spinal  Nerves  to  Reach  the  Spinal  Medulla— 
Every  Nervous  Ganglion  a  Centre  of  Nervous  Force,  Possessing  both  Sets  of  Fibres, 
or  Dilator  and  Contractor  Nerves— The  Manner  Reflex  Action  is  Produced  in  the 
Spinal  Cord  and  Medulla  Oblongata  through  Sensory  Impressions  in  the  Mucous 
Membrane  and  Cutaneous  Surface,  for  Expanding  and  Contracting  the  Vessels 
and  Maintaining:  a  Balance  in  the  Circulation — Relative  Amount  of  Nervous  Force 
sent  to  the  Viscera  through  the  Pneumogastric  and  Splanchnic  Nerves,  Dlus- 
trated  by  a  Case  of  Fracture  of  the  Fourth  and  Fifth  Cervical  Vertebrae,  Producing 


XVlll  CONTENTS. 

CHAPTER    XVIII. 

DUALISM  IX  MUSCLES  AXD  NERVOUS  FORCE. 

Nature  of  Vital  Force — Principle  in  Expansion  and  Contraction — Molecular  Changes  in 
the  Cell-Contents  Involved  in  both  Movements  ;  Illustrated  by  the  Action  in  Pro- 
toplasm and  in  Muscle  Cells  during  Contraction,  as  Seen  in  the  Field  of  the  Polar- 
izing Microscope — Dualism  in  Nervous  Force  Essential  to  the  Production  of  both 
Movements — Extraordinary  Hardness  Produced  in  the  Muscles  by  Nervous  Force — 
Explanation  Therefor— Hardness  a  Measure  of  Work— Mode  of  Demonstrating  This 
Circumstance — An  Easy  Method  of  Proving  Dualism  in  Muscles,  and  by  Implication 
Nervous  Force— Lessons  Taught  by  the  Phenomena  in  the  Leech— Ditto,  Tongue  of 
the  Frog— Ditto,  Tortoise  ;  the  Action  in  the  Head,  Neck  and  Tail  Demonstrating 
Dualism  in  Muscles  and  Nervous  Force— Ditto,  Conchifera,  for  Opening  and  Clos- 
ing the  Valves  ;  Together  with  Physiological  Experiment  Demonstrating  the  Cir- 
cumstance— Ditto,  Inferior  Maxilla  in  the  Dog,  showing  the  Masseter  and  Tem- 
poralis Muscles  are  Operated  in  the  Same  Way,  the  Mouth  being  Opened  and 
Closed  by  Means  of  Expansion  and  Contraction  in  these  Muscles — Physiological 
Experiment  upon  the  Nerves  to  the  Ciliary  Ganglion  Proving  Dualism  in  Nervous 
Force  upon  the  Nerves — The  Circumstance  Applied  to  the  Oral  Muscles  and  the 
Action  in  all  the  Sphincters,  all  of  Them  in  Common  Possessing  Circular  and 
Radiating  Muscles,  the  Same  as  the  Iris— Action  in  Erectile  Tissue  Readily  Ex- 
plained ;  Elucidated  by  the  Action  in  the  Tongue  of  the  Chameleon,  Penis,  etc. — 
The  Special  R61e  in  Nerves  with  Respect  to  Nervous  Currents,  Nervous  Centres, 
Separators  and  Delimitators  of  Electrical  Fluids  Generated  in  the  Tissues  and 
Carried  to  the  Centres  for  this  Purpose — Reasons  Therefor       ...        -        444 


Library  of  tke  New  York 
State  Medical  Association. 


CHAPTER  I. 

MECHANICS   FOE   PRODUCING    CIRCULATION  IN  AIR  AND  WATER, 

AND  FOR  CONNECTING  THEM  WITH  CIRCULATION  IN  LIVING 

ORGANISMS. 

The  Fundamental  Fact  at  the  Basis  of  Living  Organisms — Living  Organisms  Evolved 
from  Substances  Contained  in  Air  and  Water — Mode  of  Suspending  Atmospheric 
Matter  in  Space,  and  the  Principle  for  Producing  Movement  Among  the  Mole- 
cules— Experiments  of  Dufay  and  Faraday  with  Electricity  and  Magnetism — Action 
of  Gravitation  Overcome  by  Electrical  Force — Transformations  of  Heat  into  Elec- 
trical Force  for  Energizing  the  Polar  Forces  Among  the  Molecules,  the  Fundamental 
Principle  for  Producing  Circulation  in  Air  and  Water — The  Prodigious  Force  which 
is  Involved  for  Producing  Circulation  in  the  Atmosphere  and  in  Water — The 
Mechanics  Connected  Through  and  Through  by  Means  of  Electrical  Force — 
Electrical  Phenomena  in  Storms — Mode  of  Effecting  Changes  in  Pressure  for 
Increasing  Circulation  in  the  Atmosphere — The  Action  in  Water — Principle  in 
Diffusion,  Osmose  Capillarity — Organic  Connection  Subsisting  Between  Circula- 
tion in  Air  and  Water  and  in  the  Living  Organism. 

Logically  as  well  as  scientifically,  the  fundamental  fact  at 
the  basis  of  living  organisms  is  the  one  of  circulation,  since 
it  is  by  means  of  this  the  materials  are  furnished  for  elabo- 
rating structure  and  evolving  force ;  otherwise,  neither  the 
building  itself  could  have  construction,  nor  could  the 
special  vital  phenomena  be  produced,  since  they  are  all 
evolved  out  of  circulation  from  substances  which  are  brought 
into  the  organism  by  this  means,  and  which,  of  course,  would 
include  the  psychical  with  the  rest,  being  a  variety  simply, 
and  as  much  dependent  upon  circulation  as  any  of  the  others. 
In  other  words,  should  circulation  stop,  it  all  stops,  and  which 
is  a  matter  so  obvious  that  it  needs  no  argumentation. 

In  the  second  place,  the  nutritive  and  force-producing  ele- 
ments are  contained  in  air  and  water,  notably  oxygen  and 
hydrogen,  carbon  and  nitrogen,  together  with  the  alkaline 
earths  and  minerals.  Moreover,  are  incessantly  in  motion  in 
every  conceivable  direction,  both  the  fluids  themselves  and 
the  molecular  and  atomic  constituents,  so  that  it  may  be  truly 


2  ATMOSPHF.KIC   CONSTITUTION. 

said  they  are  never  still  save  when  locked  in  crystal  and 
living  structure  ;  and  even  here  a  degree  of  movement  takes 
place,  but  more  in  the  latter  than  in  the  former,  and  more  in 
animal  than  in  vegetal  structure,  while  the  freest  motion  is 
that  which  obtains  in  the  conditions  of  air  and  water  in  which 
the  molecules  and  atoms  have  the  widest  action,  the  atoms  with 
the  molecules.  Notwithstanding,  it  is  all  organized,  the  air  and 
water  with  floral  and  animal  life,  a  definite  molecular  constitu- 
tion being  maintained  throughout.  Thus,  in  the  case  of  the 
atmosphere  the  relative  proportion  of  the  constituent  gases 
remains  the  same,  or  about  77  parts  of  nitrogen,  21  of  oxy- 
gen, and  2  parts  made  up  of  watery  vapor,  carbonic  acid, 
ammonia,  etc.,  in  the  100.  This,  notwithstanding  the  enor- 
mous sources  of  disturbance  in  the  countless  flora  and  fauna, 
inclusive  of  the  chemical  reactions  taking  place  in  the  earth 
itself,  together  with  the  combustions  and  chemical  reactions 
effected  through  the  agency  of  man,  which  are  constantly 
absorbing  and  giving  out  the  gases  in  prodigious  amount, 
and  which  proves  beyond  the  shadow  of  a  doubt  the  action 
of  an  all-pervading  force  for  effecting  it,  or  for  compelling  the 
molecules  into  their  relative  positions  in  the  atmospheric 
envelope.  In  short,  that  a  principle  obtains  among  the  indi- 
vidual atoms  and  molecules  for  producing  movement  in  them, 
and  acting  in  such  wise  as  to  compel  definite  arrangements 
among  them,  to  the  end  that  the  atmosphere  and  living  struc- 
ture may  be  organized  and  a  balance  maintained  throughout, 
otherwise  is  inexplicable. 

In  the  case  of  water,  we  find  that  here  also  the  same  princi- 
ple applies  for  producing  movement  in  the  atoms  and  molecules 
for  compelling  absorption  and  solution  of  the  salts,  so  as  to 
effect  the  chemical  and  physical  constitution  that  obtains  in  this 
fluid,  and  in  order  to  maintain  a  balance  which  tends  inces- 
santly to  be  disturbed,  at  the  same  time,  producing  perpetual 
motion  throughout  in  the  mass  and  in  the  molecules  the  same 
as  in  the  air ;  using  an  expressive  term,  the  whole  seeming,  as 
it  were,  "alive."  The  ultimate  fact  in  the  deep  mystery  is 
the  one  of  motion  itself,  the  manner  it  is  produced,  with  the 
principle  it  involves;  for  life  is  evolved  by  means  of  it,  and 
may  be  characterized  as  a  metamorphosis  of  force  ;  seeing  also 


FORCE.  3 

that  when  life  terminates  the  whole  breaks  up  into  its  atomic 
and  molecular  elements  and  goes  back  whence  it  came,  and 
there  is  nothing  left. 

While  to  this,  again,  must  be  added  the  deeply  suggestive 
fact  that  organic  unity  implies  similarity  in  force.  So  that 
continuity  in  force,  which  undoubtedly  exists,  would  imply 
that  vital  force  is  but  a  variety  of  the  force  which  produces 
molecular  action  in  air  and  water,  an  organic  connection  sub- 
sisting between  them  for  producing  the  currents  in  and  out  the 
organism  in  the  measure  of  the  requirements,  otherwise  im- 
possible ;  at  once  indicating  the  nature  of  the  force  for  effect- 
ing the  nutritive  processes,  together  with  the  other  phe- 
nomena, the  whole  being  electrical  and  polar.  Thus,  in 
the  flora,  in  which  life  has  simplest  expression,  the  nutritive 
and  force-producing  elements  are  pulled  into  the  organism 
by  the  action  of  the.  polar  forces  upon  the  principle  which 
obtains  in  a  magnet,  while  for  expelling  waste  products  the 
action  is  simply  reversed.  In  the  transit  the  component  atoms 
and  molecules  are  pulled  into  position  and  locked  in  the  struc- 
tures by  the  action  of  the  polar  forces  upon  the  same  principle 
precisely  which  obtains  in  crystal  structure,  only  the  action 
is  more  complex  ;  while  the  waste  products  are  borne  out  for 
redistribution  by  action  of  the  same  forces,  distribution  with 
redistribution  being  the  eternal  order.  In  animal  life  there  is 
still  greater  complexity,  which  is  involved  in  producing  the 
characteristic  phenomena,  notably  the  multitudinous  actions 
taking  place  in  them.  At  the  same  time,  a  special  arrangement 
obtains  for  increasing  circulation  commensurate  with  this  cir- 
cumstance, and  for  producing  a  balance,  which  the  scheme  calls 
for ;  for  everything  is  evolved  out  of  circulation  from  sub- 
stances brought  into  the  organism  by  this  means,  as  before 
remarked. 

Finally,  since  all  this  motion  involves  mechanical  work,  it 
would  seem  both  reasonable  and  natural  that  the  principle 
underlying  it  should  be  susceptible  of  formulation  and  distinct 
mental  presentation  A  force  in  such  universal  and  constant 
action  should  surely  be  detectable,  if  only  we  go  about  it  in 
the  proper  way. 

In  the  first  place,  let  us  take  up  the  action  in  the  atmosphere. 


C  FORCE. 

This  matter  settled,  and  there  will  be  no  difficulty  in  account- 
ing for  circulation  in  living  organisms,  which  is  but  an  exten- 
sion of  the  same  principle  in  mechanics,  as  we  shall  very 
plainly  demonstrate  in  the  succeeding  pages,  the  matter  being 
one  of  variety  simply. 

Concerning  the  Force  for  Effecting  the  Suspension  of  Atmos- 
pheric Matter  in  Space  and  for  Producing  incessant 
Motion  in  it  against  the  Force  of  Gravitation  : 

Commencing  at  the  bottom,  then,  so  to  speak,  and  dealing  only 
with  the  forces  in  Nature,  the  first  question  which  presents  itself 
to  the  thoughtful  student  concerns  the  mechanics  for  effecting 
the  suspension  of  atmospheric  matter  in  space  and  the  pro- 
duction of  incessant  motion  in  it  against  the  force  of  gram 
tation,  which  tends  to  pull  it  to  the  earth  in  a  compact  mass. 
In  short,  why  should  there  be  an  atmosphere  at  all  upon  the 
globe  ?  and  how  is  motion  produced  in  it  % — facts  which  are 
certainly  needing  explanation. 

The  atmospheric  molecules  do  not  touch  each  other,  as  in  the 
case  of  a  solid  ;  on  the  contrary,  they  are  far  removed  from 
each  other,  so  that  even  at  the  earth's  surface,  where  density 
is  greatest,  the  atmosphere  would  require  to  be  compressed 
fully  700  times  to  reduce  it  to  the  density  of  a  liquid,  while 
the  molecules  recede  farther  and  farther  from  each  other  with 
the  distance  from  the  earth  till  the  outermost  limits  are  reached 
(Fig.  1).  In  point  of  fact,  the  molecules  are  relatively  as  far 
apart  in  proportion  to  size  as  the  very  stellar  masses  them- 
selves :  furthermore,  when  forcibly  compressed,  react  with 
equal  force,  at  once  springing  back  to  the  original  distances 
the  moment  pressure  is  relieved,  proving  thereby  inter-molecu- 
lar action  from  the  presence  of  a  force  radiating  from  molecule 
to  molecule,  otherwise  is  inexplicable.  Now,  then,  this  fact 
being  indisputable,  the  question  which  naturally  presents 
concerns  the  nature  of  this  force.  What  is  it  ?  This  must  be 
established  before  progress  can  be  made  in  the  mechanics  of 
circulation,  forming,  as  it  were,  the  basis  of  the  mechanics, 
and  by  means  of  which  the  nutritive  and  force-producing 
elements  are   set   in   motion   for  elaborating   structure   and 


ATMOSPHEEIC   MAI  TEE. 


evolving  force,  the  whole  connecting  through  and  through,  the 
matter  necessarily  involving  continuity  in  force  in  order  to 
effect  these  results. 


•  •  • 


_  o 

Jo 


°     o     0  o 

o      ^         °a9       ° 

0  o    o°    o"j    ° 

1  °    °     o     o     o    0 

O     o     O        O     q 

>o  °     o   °     o      ° 


O   o 


,  ©  °  °< 

o   o   o 


o  o 


o    o 
o  o 


_o  o   0 
o    ° 


°  o' 


o  o 


\j  _  O    O    q     o        o 

,0o°00oVo0oo00°o0°oOo°oO°Ooo:jo-0 
°o°°oro0o0=V0Oo°co^o„o0°,oo0o0 

0--0-«--V.°-,0o 

000  °  °o"o0°0o0o0!;o  °  °oo" 


Fig.  1. — An  Ideal  Section  of  the  Atmosphere,  showing  the  Suspension  of  this  Matter  in 
Space  against  the  Force  of  Gravitation,  a,  a,  Molecules  of  Nitrogen,  Oxygen,  Car- 
bonic  Acid,  etc. ;  b,  b.  Aqueous  Molecules. 

In  this  we  are  materially  assisted  by  the  celebrated  experi- 
ment of  Dufay  for  proving  the  existence  of  two  electricities, 


6  POLAll   FOKCES. 

or  positive  and  negative,  which  is  plain  enough.  For  ex- 
ample, he  discovered,  by  means  of  a  glass  rod  excited  by 
rapid  friction,  that  a  gold  or  silver  leaf,  when  liberated 
near  it,  is  both  attracted  and  repelled  by  it.  At  first, 
is  powerfully  attracted,  dashing  impetuously  toward  it,  and, 
suddenly  stopping,  dashes  in  the  opposite  direction,  when  it 
again  stops  suddenly,  remaining  stationary  from  two  to  three 
inches  from  the  rod,  suspended  in  the  air  (Fig.  2).  And  if  one 
so  mind,  he  may  chase  it  round  and  round  the  room  for  hours 
without  permitting  it  to  fall  to  the  ground.  The  knob  of  a 
charged  Leyden  jar,  or  the  conductor  of  an  electrical  machine 
in  action,  has  the  same  effect ;  and  which  proves  the  action 
is  due  to  electricity.  The  use  Dufay  made  of  this  experi- 
ment was  to  demonstrate  the  existence  of  two  electricities, 
or  positive  and  negative,  whereby  substances  are  attracted 
and  repelled  from  each  other,  according  to  whether  they  are 
charged  with  opposite  or  the  same  kind  of  electricity. 

The  attraction  in  the  first  instance  was  due  to  opposite  elec- 
tricities, while  the  subsequent  repulsion  was  due  to  the  passing 
over  of  some  of  the  electricity  from  the  more  highly  electrified 
glass  to  the  metallic  leaflet,  whereby  their  properties  were 
assimilated  ;  hence  this  repulsion.  The  point  where  it  remains 
stationary  represents,  of  course,  equilibrium  in  the  polar 
forces.  But,  then,  this  describes  only  a  part  of  the  phenomena 
embraced  in  this  experiment,  which  admits  of  much  wider 
application  than  proving  the  existence  of  two  electricities 
simply  ;  notably,  the  mechanical  worJc  which  is  involved  in 
sustaining  matter  in  space,  and  producing  movement  in  it 
against  the  force  of  gravitation,  which  tends  to  pull  it  to  the 
earth.  Thus  two  great  and  important  elements  in  this  elec- 
trical experiment  are  entirely  overlooked:  1.  The  fact  that 
matter  may  be  held  up  and  sustained  in  space  by  electrical 
force  simply.  2.  That  motion  can  be  produced  in  it  by  a 
change  in  polarity  effected  through  electrical  induction — facts 
which  are  incontrovertible. 

Now,  then,  proceeding  cautiously,  feeling  the  ground,  as  it 
were,  to  make  sure  of  its  firmness :  Is  there  any  other  force 
in  Nature  for  thus  suspending  matter  in  space  and  producing 
movement  in  it  against  the  force  of  gravitation,  save  electricity 


ELECTRICAL   FORCE.  7 

and  the  interaction  of  the  polar  forces  which  electricity  serves 
to  intensify  \  Not  that  we  know  of.  No,  not  any  force  what- 
ever may  be  spoken  of  in  this  connection,  save  electricity 
alone.  And  making  deduction  from  this  upon  the  mechanics 
in  the  atmosphere,  it  follows  that  electricity  is  the  force  we 
are  in  search  of  for  effecting  the  suspension  of  atmospheric 
matter  in  space,  and  for  producing  movement  in  it,  no  other 
force  applying  for  the  purpose.  This  fact  conceded,  the 
next  question  concerns  the  source  of  electrical  supply, 
together  with  the  manner  of  its  action,  which  is  also 
obvious.     Thus  the  sun  functions  as  the  source  of  electrical 


Fig.   2. — Showing  Matter  Suspended  in  Space  by  interaction  of  the  Polar  Forces, 
intensified  by  electrical  induction. 

supply,  by  means  of  which  the  Leyden  jar  in  the  earth  itself 
is  kept  constantly  charged  with  electricity,  whence  it  radiates 
through  the  atmosphere  by  the  principle  of  electrical  induc- 
tion for  effecting  suspension  of  this  matter  in  space  (Fig.  1), 
and  for  producing  movement  in  it.  In  other  words,  heat, 
impinging  against  the  earth,  undergoes  transformation  into 
electrical  force,  which  is  the  motor  in  the  mechanics.  That 
heat  per  se  is  not  the  motor,  is  at  once  made  obvious  by 
the  fact  that  rarefaction  in  the  atmosphere  increases  with 
the  cold  and  the  distance  from  the  earth  (Fig.  1),    being 


8  SOLAR   FORCE. 

greatest  at  the  outermost  limits  in  the  atmosphere  where  cold  is 
enormous,  whereas  the  very  opposite  should  obtain  by  the  laws 
of  radiation,  the  surface  being  the  first  chilled,  producing  con- 
densation— an  increase  in  density.     In  short,  producing  a  com- 
pact mass.     In  lieu  of  this,  however,  the  inter-molecular  dis- 
tances are  vastly  increased,  showing  conclusively  that  heat  is 
not  the  force  for  effecting  this  action.     In  point  of  fact,  the 
atmosphere  is  nearly  transparent  to  heat,  and  but  for  the 
aqueous  vapor  suspended  in  it,  nearly  the  whole  of  the  solar 
beam  would  reach  the  earth.     And  allowing  for  this  circum- 
stance, Professor  Tyndall  estimates  the  invisible  rays  or  the 
heat  portion  of  the  solar  spectrum  to  approximate  that  in  the 
spectrum  of  electric  light,  or  nearly  eight  times  the  visible  rays 
(Fig.  3),  which  will  give  some  idea  of  the  enormous  role  which 
heat  performs  in  terrestrial  mechanics.     Furthermore,  it  is  illu- 
sory to  imagine  that   all  this  heat  radiates  into  space,  since 
a  prodigious  amount  undergoes  transformation  into  mechanical 
work  and  electricity,  which  is  the  dynamic  force  for  produc- 
ing molecular  action  by  increasing   the  polar  forces  in  the 
molecules,  and  which  applies  to  living  as  well  as  to  non-living 
matter.     Perhaps  we  might  illustrate  this  circumstance  better 
by  drawing  from  the  early  experiences  in  animal  temperature. 
For  example,  after  Lavoisier  had  established  the   fact  that 
animal  temperature  is  the  result  of  oxidation,  upon  the  same 
principle  precisely  as  obtains  in  a  grate  and  the  burning  of 
coal,  physicists  made  great  efforts  to  determine  the  equality 
between  the  theoretical  heat  as  indicated  by  the  amount  of 
carbonic  acid  discharged  and  the  quantity  of  heat  furnished 
\yf  the  animal,  and  which,  of  course,  was  illusory,  for  it  made 
no  allowance  for  the  large  amount  which  disappears  by  con- 
versions into  mechanical  work  and  electricity  for  producing  the 
multitudinous  actions   taking  place  in  them.     And  making 
the  same  deduction  for  the  solar  beam  and  the  mechanics  for 
circulating  air  and  water,  inclusive  of  the  actions  in  living 
organisms,  it  at  once  becomes  obvious  that  a  vast  amount  of 
heat  must  necessarily  disappear  in  the  transformations. 

Then,  again,  one-half  only  of  the  earth  presents  to  the  sun 
at  a  given  time,  and  as  electrical  force  may  make  the  circuit 
of  the  earth  in   one-tenth  of  a  second   (as  the  solar  beam 


HEAT. 


9 


travels  nearly  12,000,000  miles  in  a  minute),  it  follows  that  this 
should  call  for  corresponding  increase  in  the  transformations 
of  heat  for  energizing  the  actions  in  the  dark  portions  of  the 


earth,  and  which  wonld  reduce  in  proportion  the  radiation 
into  space.     Hence,  we  must  conclude  that  the  greater  pro- 


10  THE   FORCE   IX   AERIAL   MECHANICS. 

portion  of  the  heat  in  the  solar  beam  is  utilized  in  the  terres- 
trial mechanics,  since  every  movement  calls  for  corresponding 
expenditure  of  force.  Finally,  these  conversions  of  heat  into 
mechanical  work  and  electricity  are  facts  which  are  uni- 
versally conceded  in  science,  being  fully  established  and  incon- 
trovertible. Impinging  against  the  earth,  then,  this  invisible 
portion  of  the  solar  beam  rebounds  as  electrical  force  through 
the  atmosphere,  driving  and  scattering  the  molecules  in  every 
possible  direction.  In  fine,  the  impact  of  the  solar  beam 
against  the  earth  fractures  and  scatters  the  atmospheric  en- 
velope in  accordance  with  well-known  physical  laws,  or  as  a 
hammer  does  a  piece  of  glass,  the  force  radiating  from  the 
point  of  impact  and  driving  the  outermost  fragments  further- 
est  apart.  Hence,  rarefaction  in  the  atmosphere  increases 
with  the  distance  from  the  earth.  This  fact,  then,  would  ex- 
plain that  circumstance,  and  no  other  could.  And  each  mol- 
ecule of  the  constituent  gases  being  charged  by  the  Leyden 
jar  in  the  earth  itself,  becomes  in  its  turn  a  centre  of  force 
which  is  readily  affected  by  the  polar  forces  in  the  earth,  so 
that  the  conditions  obtain  for  producing  movement  in  them, 
the  whole  matter  depending  upon  changes  in  polarity.  It  is 
needless  to  add  that  no  amount  of  cold  could  affect  this  action, 
which  is  precisely  what  the  circumstances  in  aerial  mechanics 
calls  for,  since  incessant  motion  pervades  it  all. 

Some  idea  of  the  enormous  force  which  is  involved  in 
aerial  mechanics  may  be  had  from  the  atomic  weight  of  the 
constituent  gases — notably  nitrogen,  L4  ;  oxygen,  16;  aqueous 
vapor,  18 ;  carbonic  acid,  44,  or  nearly  two  and  one-half 
times  greater  than  water ;  while  the  pressure  of  the  whole 
is  equal  to  33  feet  of  water  spread  over  the  entire  surface  of 
the  globe.  And  yet  it  is  all  lifted  up  in  space  and  borne 
hither,  thither,  in  every  possible  direction,  with  the  utmost 
ease  and  celerity,  under  the  action  of  electrical  force,  as  illus- 
trated by  the  rubbed  glass  and  silver  leaf,  only  that  in  the 
present  instance  the  action  is  automatic.  And  proceeding 
upon  this  line  of  investigation,  we  will  now  take  up  some  of 
the  actions  in  the  atmosphere,  both  as  regards  the  currents 
which  exist  in  it  and  the  inter-molecular  movements  for 
maintaining  its  organic  constitution,  together  with  the  cur- 


POLAR   ACTION   IN   THE   ATMOSPHERE. 


11 


rents  which  set  in  and  out  the  flora  and  fauna,  all  which 
respond  readily  to  treatment,  and  while  the  action  is  complex, 
it  unravels  easily  by  the  key  furnished  in  the  polar  forces 

Given  that  atmospheric  matter  is  held  in  space  by  the  inter- 
action of  the  polar  forces  (and  which  must  be  conceded,  since 
there  must  be  a  force  for  overcoming  gravitation),  it  follows 
that  the  various  movements  which  pervade  the  atmosphere 
must  also  be  due  to  this  circumstance  ;  or,  in  other  words,  they 
flow  out  of  the  polar  forces  as  water  from  a  fountain,  which 
cannot  rise  higher  than  its  source. 

And  the  molecules,  thus  resting  upon  the  polar  forces,  as  the 

A 


V? 


Wi. 


%,;■. -■.  a>- 


Fig.  4. 

Jloor  of  support  against  the  action  of  gravitation,  it  follows 
that  any  change  in  polarity  must  necessarily  affect  the  relations 
of  the  molecules.  It  could  not  be  otherwise,  in  the  very  nature 
of  things.  No  w,  then,  taking  this  as  the  basis  of  the  mechanics 
(and  everything  must  have  a  basis),  we  will  begin  with  the 
larger  movements  first,  proceeding  thence  to  the  inter-molecular 
for  maintaining  organic  unity,  thence  to  the  actions  in  living- 
organisms. 

Concerning  the  Currents  in  the  Atmosphere : 
In  this  investigation  we  are  materially  assisted  by  the  re- 


12 


polar  actio:;. 


searches  of  Faraday,  by  means  of   which  the  action  in  the 
atmosphere  is  brought  clearly  into  view.     For  example,  he 


Fig.  5. — Magnetic  Lines  of  Force. 
From  a  Photograph  by  Professor  Mayer. 


POLAR   ACTION. 


13 


discovered,  by  placing  a  sheet  of  smooth  paper  or  glass  over 
a  bar-magnet  and  showering  fine  iron  filings  upon  it,*  that 
they  would  arrange  themselves  in  systematic  order  and  in 
beautiful  curved  lines,  extending  from  pole  to  pole  (Figs.  4, 
5,  6).  Now,  by  applying  this  circumstance  to  the  earth  itself 
and  the  actions  taking  place  in  air  and  water,  notably  the 
currents  setting  in  and  out  of  the  poles,  together  with  the 
formation  of  the  cloud  strata,  are  at  once  made  intelli- 
gible.     If  the  earth  itself  be  a  prodigious  magnet  (which. 


Fig.  6. — Showing  the  "Lines  of  Force"  extending  from  Pole  to  Pole. 


no  one  should  deny,  since  it  could  not  otherwise  be  held  in 
space,  nor  motion  be  produced  in  it),  and  atmospheric  matter 
is  suspended  in  space  by  inter-action  of  the  polar  forces,  as 
alleged,  it  follows  that  similar  arrangements  should  obtain 
among,  the  molecules  as  in  the  iron  filings,  the  same  law  apply- 
ing for  both.  Furthermore,  that  there  are  such  k ;  lines  of 
force"  stretching  their  long  arms  between  the  terrestrial  poles, 

*  The  iron  filings  being  very  heavy,  and  consequently  not  free  to  move,  in 
order  to  facilitate  the  action  "the  sheet  is  gently  tapped  from  time  to  time 
so  as  to  release  them  for  a  moment,  and  enable  them  to  follow  their  tend- 
encies." 


14  TERRESTRIAL   MAGNETISM. 

is  fully  established  by  the  action  taking  place  in  the  magnetic 
needle,  which,  the  moment  it  is  suspended  and  free  to  move, 
trembles,  oscillates  and  rocks  from  side  to  side,  nor  pauses  till 
it  is  brought  into  correspondence  with  the  magnetic  meridian, 
one  end  pointing  north,  the  other  south,  while  every  atom  and 
molecule  feels  the  pull  in  these  polar  forces;  and  should  they  be 
free  to  move  as  the  molecules  in  air  and  water,  can  it  be  doubted 
for  a  single  moment  but  that  they  would  be  pulled  to  their 
respective  poles,  the  positive  going  one  way  and  the  negative 
the  other  ;  since  it  is  reasonable  to  assume  the  forces  which 
coerce  them  in  mass  should  be  equally  effective  upon  the  indi- 
vidual molecules  ?  This,  then,  would  explain  that  matter,  and 
n  >  other  could.  Faraday  !  thou  incomparable  !  we  thank  thee  ! 
This  experiment  of  yours  with  the  magnet  puts  iron  through 
the  logic.  The  cloud  strata,  the  currents  in  the  air,  and  the 
currents  in  the  ocean  are  explicable  by  polar  action  only, 
while  it  all  represents  forms  of  conversion  of  force.  Passing 
rapidly  over  the  actions,  for  the  sake  of  brevity,  it  would 
explain  the  following  phenomena,  otherwise  inexplicable  : 

1st.  Why  the  storm-centre  does  not  follow  the  isothermal 
lines,  pursuing  an  erratic  course,  and  going  in  every  possible 
direction. 

2d.  The  occurrence  of  violent  storms  in  the  polar  regions,  and 
their  greater  frequency  in  the  temperate  zones  during  winter. 

3d.  It  would  account  for  whirlwinds,  in  which  the  air  pur- 
sues a  violent  rotatory  motion,  since  this  is  one  of  the  phe- 
nomena which  accompany  electrical  excitement.  Moreover,  a 
miniature  whirlwind  is  readily  reproduced.*  And  since  elec- 
tricity increases  magnetism,  this  circumstance  would  explain 
the  cohesive  power,  so  to  speak,  among  the  molecules,  which 
imparts  corresponding  firmness  and  strength  to  the  atmosphere, 

:- A  curious  "  object-lesson  "  is  that  by  which  Professor  Douglass,  of  the 
Michigan  State  University,  shows  his  classes  the  operation  of  a  cycl&ne.  He 
suspends  by  silken  cords  a  large  copper  plate,  which  is  heavily  charged  with 
electricity.  The  electricity  hangs  down  underneath  the  plate  like  a  bag,  and 
is  rendered  visible  by  the  use  of  arsenious  acid  gas,  which  gives  it  a  green  color. 
The  formation  is  a  miniature  cyclone  as  perfect  as  any  started  in  the  clouds. 
It  is  funnel-shaped,  and  whirls  around  rapidly.  Passing  this  plate  over  a  table, 
the  cyclone  snatches  up  copper  cents,  pith  balls  and  other  objects,  and  scatters 
them  on  all  sides. 


ATMOSPHEEIC    MAGNETISM.  15 

whereby  solid  masses  of  great  weight  are  caught  up  and  trans- 
ported to  great  distances  in  the  funnel  of  the  whirl  wind. 

4th.  It  would  explain  why  thunder-storms  are  accompanied 
by  vivid  electrical  phenomena  and  the  fall  of  the  thunder-bolt. 
The  electrical  force  which  suspends  water  in  the  atmosphere 
being  withdrawn,  an  amount  of  electricity  representing  this 
circumstance  passes  at  once  to  the  earth,  and  striking  the  object 
most  highly  charged  at  the  time  with  opposite  electricity,  dif- 
fuses itself  through  the  earth  for  producing  equilibrium.  In 
fine,  water  falls  because  the  thunder-bolt  falls,  the  one  produc- 
ing the  other. 

5th.  It  would  account  for  the  cloud  corona  resting  over 
islands  in  the  sea,  and  why  vegetation  should  milk  the  clouds. 
All  these  circumstances,  then,  have  ready  explanation  by  this 
mechanics,  and  they  can  be  explained  in  no  other  way.  But 
we  are  by  no  means  done  yet;  a  multitude  of  phenomena  other- 
wise inexplicable  remain  for  mention. 

The  following  forcible  excerpt  will  be  appreciated : 
' '  The  experiments  of  Faraday  show  that  the  magnetism  of 
the  air  changes  with  temperature  ;  that  it  is  least  near  the 
equator,  and  greatest  at  the  poles  of  maximum  cold ;  that  it 
varies  with  the  seasons,  and  changes  night  and  day  ;  nay,  the 
atmosphere  has  regular  variations  in  its  electrical  conditions, 
expressed  daily  at  stated  hours  of  maximum  and  minimum 
tension  Coincident  with  this,  and  in  all  parts  of  the  world, 
but  especially  in  sub-tropical  latitudes,  the  barometer  also  has 
its  maxima  and  minima  readings  for  the  day.  So  also,  and 
-at  the  same  hours,  the  needle  attains  the  maxima  and  minima 
of  its  diurnal  variations.  Without  other  time-piece,  the  hour 
of  the  daymay.be  told  by  these  maxima  and  minima,  each 
group  of  which  occurs  twice  a  day  and  at  six-hour  intervals. 
These  invisible  ebbings  and  Sowings — the  diurnal  change  in 
the  electrical  tension — the  diurnal  variation  of  the  needle,  and 
the  diurnal  rising  and  falling  of  the  barometer,  follow  each 
other  as  closely  and  as  surely,  if  not  quite  as  regularly,  as 
night  the  day.  Any  cause  which  produces  changes  in  at- 
mospheric pressure  invariably  puts  it  in  motion,  giving  rise  to 
gentle  airs  or  furious  gales,  according  to  degree  ;  and  here,  at 
least,  we  have  a  relation  between  the  movements  in  the  air  and 


](')  ELECTRICITY   PRODUCING   CIRCULATION 

the  movements  of  the  needle  so  close  that  it  is  difficult  to  say 
which  is  cause,  which  effect,  or  whether  the  two  be  not  the 
effects  of  a  common  cause  "* 

The  mode  of  increasing  the  action  in  the  air  is  by  changing 
pressure,  notably  by  developing  areas  of  low  pressure  with 
an  increase  in  pressure  in  adjacent  localities,  whereby  fluid 
equilibrium  is  invoked,  the  fluids  rushing  in  to  equalize  press- 
ure. And  the  greater  the  difference  in  the  low  and  high 
pressures,  the  more  violent  the  storm,  while  the  whole  is  due 
to  electrical  disturbance.  Taking  the  area  of  low  pressure  as 
the  storm  centre,  we  must  regard  this  as  due  to  a  flood  of 
electrical  force  which,  by  increasing  polarity  among  the  mole- 
cules causes  them  to  recede  farther  and  farther  from  each 
other,  and  in  proportion  producing  aerial  expansion  with 
corresponding  fall  in  pressure,  while  the  adjacent  fluids  flow 
into  the  locality  till  pressure  is  again  uniform.  So,  then,  for 
producing  the  atmosphere  itself,  we  need  the  action  of  elec- 
tricity; and  for  producing  currents  in  it,  we  need  the  action  of 
electricity;  finally,  for  energizing  the  movements  for  producing 
the  rapid  currents,  we  need  the  action  of  electricity.  The 
curious  circumstance  that  pressure  falls  when  the  air  is  satu- 
rated with  moisture,  is  to  be  explained  in  the  same  way. 

Thus  for  effecting  evaporation,  electrical  force  is,  so  to  speak, 
piled  up  in  the  aqueous  molecules  which,  by  riving  them  widely 
asunder,  floats  them  off  into  the  atmosphere,  but  in  place  of 
increasing  density,  pressure  falls  in  correspondence  with  the 
degree  in  saturation  by  reason  of  the  fact  that  the  aqueous 
molecules,  by  acting  as  centres  of  electrical  force,  react  upon 
the  molecules  of  the  constituent  gases,  causing  them  in  turn 
to  recede  farther  and  farther  from  each  othei\  so  that  expan- 
sion and  low  pressure  are  made  inevitable.  Thus  we  have 
the  conditions  for  producing  expansion  in  the  atmosphere,  for 
increasing  circulation  correspondingly  inherent  in  the  very 
mechanics  of  evaporation,  and  the  wonderful  utilitarian 
methods  in  Nature,  have  striking  illustration,  the  matter  being 
also  self-adjusting.  In  this  manner  the  flora  and  fauna  are 
fed  out  of  the  seas  and  the  earth  is  watered.     Finally,  this 

*  The  Physical  Geography  of  the  Sea,  pp.  152,  153.    Maury. 


CHANGES   IN    PRESSURE.  17 

mode  of  increasing  circulation  in  the  air  by  rhythmical 
changes  in  pressure  effected  through  the  action  of  electrical 
force  extends  as  well  into  living  organisms  ;  notably  animals, 
in  which  it  performs  an  enormous  role,  as  indicated  in  the 
rhythmical  expansions  and  contractions  taking  place  in  the 
organs  for  pumping  the  fluids  through  the  structures  Gam- 
in snsurate  with  the  force  which  is  expended  in  them.  Other- 
Avise  the  universal  pumping  actions  going  on  in  the  body 
would  be  meaningless.  This  en  passant,  as  we  would  not 
anticipate. 

Granted  that  the  atmospheric  molecules  are  suspended  in 
space  by  electrical  induction  propagated  from  the  earth,  and 
moving  hither  and  thither  by  the  action  of  the  polar  forces, 
and  the  whole  is  at  once  made  intelligible,  together  with  the 
currents  setting  in  and  out  of  the  poles,  the  culminating  points 
of  polar  force  in  the  earth,  while  it  all  forms,  so  to  speak,  a 
magnetic  sea  in  incessant  motion  under  the  action  of  the  polar 
forces,  which  determine  all  the  movements.  And  since  rhyth- 
mical changes  in  polarity  inhere  in  polar  action,  it  follows 
that  incessant  movement  is  made  inevitable.  Then,  again,  the 
unequal  distribution  of  solar  heat  produced  by  the  spherical 
form  of  the  earth,  which  presents  only  a  moiety  of  the  surface 
at  a  time  to  the  solar  beam,  together  with  the  rotatory  motion 
upon  its  axis,  should  tend  to  increase  the  action,  not  only  as 
Ibetween  the  equator  and  the  poles,  and  opposite  sides  of  the 
earth,  but  also  the  XDerpendicular  movements  as  well,  the 
atmosphere  and  earth  possessing  opposite  polarities,  the  latter 
being  negative,  the  former  positive.  And  in  this  connection  it 
may  not  be  amiss  to  call  to  mind  the  important  fact,  equally 
suggestive,  of  the  air  circulating  in  the  very  crust  of  the  earth 
itself,  as  also  in  the  waters,  since  it  forms  an  essential  part  in 
the  economy  of  Nature  and  is  needing  explanation ;  and  were 
polar  force  eliminated  from  these  actions,  it  would  be  utterly 
impossible  to  explain  them.  For  example,  the  metals  are 
nearly  all  oxidized,  and  numerous  salts  are  formed,  while  sub- 
terranean waters  are  highly  charged  with  the  gases,  notably  car- 
bonic acid  ;  and  making  due  allowance  for  the  action  of  water, 
these  results,  in  the  main,  are  undoubtedly  due  to  the  action 
of  the  atmosphere,   which  is  the  source  of  the  free  oxygen 


18  AIR   IN   WATER. 

and  carbonic  acid;  while  in  the  ocean  itself  air  passes  in  a 
constant  stream  to  the  very  floor  for  evolving  and  sustaining 
the  life  which  is  produced  in  this  locality,  otherwise  impossi- 
ble. A  circumstance  which  is  also  deeply  interesting,  since  it 
proves  incontrovertibly  the  existence  of  interstices  in  water ; 
and  being  nearly  incompressible,  it  follows  that  water  is  formed 
of  globular  or  spheroidal  bodies  in  actual  contact,  but  leaving 
interstices,  as  a  matter  of  course,  and  similar  to  what  occurs 
when  shot  are  heaped  together,  through  which  fine  sand  is 
readily  sifted  to  the  bottom  by  means  of  agitation,  gravitation 
compelling  this  circumstance.  But  the  simile  fails  at  this 
point,  however,  for  the  air  goes  down  to  the  floor  and  returns 
in  a  constant  stream,  otherwise  a  balance  in  the  gases  could 
not  bs  maintained.  From  the  very  nature  of  the  mechanics 
this  circulation  is  difficult.  And  here  comes  in  the  great  utility 
of  the  "tides"  for  effecting  rairid  admixture  of  the  atmos- 
pheric gases  in  the  water,  and  not  a  wave  occurs  in  the  ocean 
but  contributes  its  part  for  maintaining  equilibrium  in  the 
oceanic  gases,  while  the  grand  scheme  in  Nature  receives  ad- 
ditional emphasis  ;  at  the  same  time,  the  very  principle  in 
tidal  movement  shows  incontrovertibly  that  water  is  dominated 
by  polar  force,  the  all-pervading  principle  in  Nature  for  main- 
taining continuity  in  force.  In  proof  of  this,  we  have  only  to 
look  from  the  actions  in  the  mass  to  those  taking  place  among 
the  molecules,  of  which  water  furnishes  eloquent  example. 

The  Actions  Talcing  Place  in   Water : 

The  prodigious  force  which  disappears  when  oxygen  and  hy- 
drogen combine  in  the  form  of  water  (H2  0)*  prepares  the  mind 
for  its  marvelous  performances.  This  substance  is  never  still, 
the  nearest  approach  being  when  locked  in  crystal  structure, 
while  perpetual  motion  is  the  eternal  order  ;  going  this  way, 
that  way,  and  in  every  conceivable  direction,  under  action   of 

*  Faraday  estimated  the  electrical  force  involved  in  the  decomposition  of  a 
single  grain  of  water  to  be  equivalent  to  800,000  discharges  of  his  large  Leyden 
bat  ery  in  the  Royal  Institution.  Weber  and  Kohlrausch  estimate  the  quantity 
of  electricity  associated  with  one  milligramme  or  the  g'gth  of  a  grain  of  hydrogen 
in  water,  if  diffused  over  a  cloud  1,000  metres  above  the  earth,  would  exert  upon 
an  equal  quantity  of  the  opposite  electricity  at  the  earth's  surface  an  attrac- 
tive force  of  2.268,000  kilogrammes,  or  about  5.000.000  foot-pounds. 


POLAE  ACTION   IN   WATEE.  19 

the  polar  forces  and  to  which  nothing  is  comparable  save  the 
actions  taking  place  in  animal  organisms,  and  in  which  the 
deeply  suggestive  fact  is  disclosed  that  it  forms  fully  three- 
fourths  of  the  structures,  the  gray  matter  of  the  brain  being 
80. y  decimals  water  ;  while  the  nutritive  and  force-producing 
elements  are  suspended  in  its  midst,  and  transported  through. 
the  channels  of  the  circulation  by  this  means.  Mirabile  dictu  I 
Mirabile  !  Mirabile  !  Rising  up  from  the  ocean  in  enormous 
volumes  from  the  action  of  electrical  force,  it  swims  away  in  the 
magnetic  sea,  and  descending  to  earth  from  a  change  in  polari- 
ty or  abstraction  of  force,  it  feeds  the  flora  and  fauna,  the 
excess  flowing  off  the  water- sheds  back  again  into  the  ocean, 
and  so  making  the  circuit  of  the  globe  in  this  manner  under 
action  of  the  special  polar  forces  which  apply  ;  but  also  rising 
and  falling  continually  from  the  earth  at  the  same  time,  while 
heat  serves  for  energizing  the  movements  by  increasing  polarity 
among  the  molecules,  as  before  remarked ;  otherwise  is  in- 
explicable. And  sailing  in  the  great  round  of  its  circulation  in 
the  air,  at  times  spreading  out  so  as  to  become  invisible,  but  com- 
ing together  again  and  again  to  form  the  cloud  strata  and  the 
rain  from  responsive  changes  in  polarity,  it  is  at  once  perceived 
that  the  whole  mechanics  connects  through  and  through  by 
means  of  this  all-pervading  principle ;  while  electricity 
is  the  dynamic  force  for  effecting  the  changes  in  polari- 
ty. Take  another  example.  At  a  temperature  of  2i2 
degrees  Fahr.,  water  expands  1,700  times  its  volume,  while 
J, 000  degrees  of  temperature  disappear.  Bat  how  ex- 
pand %  and  what  form  of  force  does  temperature  assume  % 
We  have  seen  that  the  only  means  for  suspending  matter 
in  space,  and  for  producing  movement  in  it,  is  by  electrical 
induction  effected  through  the  action  of  the  polar  forces 
which  electricity  intensifies,  and  by  applying  this  circum- 
stance to  evaporation  and  the  action  taking  place  in  boiling 
water,  the  phenomena  are  at  once  made  intelligible.  Thus 
in  the  case  of  boiling  water,  as  in  simple  evaporation,  heat 
undergoes  metamorphosis  into  electrical  force,  which  becomes 
piled  up  as  it  were,  in  the  molecules,  riving  them  farther  and 
farther  apart,  and  floating  them  up  in  the  magnetic  sea,  the 
greater  density  in  the  air  also  serving  to   expedite  the  move- 


20  POLAR   ACTION    IN    DIFFUSION. 

merit ;  since  no  other  force  applies  for  suspending  matter  in 
space  ami  producing  movement  in  it  save  electrical  force,  into 
which  heat  is  transformed  or  converted. 

And  thus  supported  by  electric  force,  it  follows  that  when 
this  is  withdrawn,  as  lnrppens  when  clouds  of  opposite  polari- 
ties come  in  contact,  the  water  must  again  condense  itself 
and  fall  to  the  earth  from  the  action  of  gravitation.  Hence 
the  electrical  phenomena  which  accompany  rain.  All  which 
is  readily  intelligible. 

Diffusion  :  Take  the  mechanics  in  diffusion  as  further  illus- 
tration of  this  principle  in  molecular  action,  and  see  how 
readily  this  also  responds  to  treatment. 

It  is  seen  to  great  advantage  in  dissolving  sugar.  For  ex- 
ample, if  a  lump  of  loaf-sugar  be  held  against  the  side  of  a 
glass  of  water  and  suffered  to  dissolve,  a  current  of  fine  saccha- 
rine matter,  in  form  of  a  white  cloud,  is  seen  to  issue  from 
every  portion  of  the  mass  and  circle  round  and  round  the  limits 
in  the  glass,  going  first  toward  the  bottom,  seemingly  from  the 
action  of  gravitation,  but  crossing  over  and  ascending  on  the 
opposite  side  till  near  the  surface  waters,  comes  back  again  to 
the  mass,  then  down  again  toward  the  bottom,  as  before,  in  a 
continuous  circuit,  nor  pauses  till  all  the  sugar  is  dissolved  or 
the  water  becomes  fully  saturated. 

Now,  then,  if  polar  force  were  eliminated,  how  would  it  be 
possible  to  explain  this  action?  Moreover,  should  some  ar- 
rangement obtain  for  removing  the  sugar  as  fast  as  introduced, 
no  reason  exists  why  it  should  not  be  perpetual,  or  why  it 
should  not  belt  the  globe  itself,  had  the  water  such  extension. 

Here,  then,  we  have  circulation  produced  by  polar  force  sim- 
ply, while  heat  serves  for  energizing  it  by  increasing  polarity 
among  the  molecules,  and  in  this  manner  disappearing  in 
mechanical  work.  In  short,  diffusion,  whether  it  relate  to 
atmosphere  or  water,  or  to  substances  undergoing  solution, 
is  universally  the  same,  being  due  to  the  action  of  the  polar 
forces,  and  which  is  fully  corroborated  by  the  fact  that  heat 
serves  to  energize  the  action  in  all  of  them,  and  showing  the 
principle  is  all-pervading. 

Osmose  and  Capillarity :  Continuing  the  inquiry,  take 
.osmose  and  capillarity,  circumstances  so  much  relied  upon  for 


POLAR  ACTION  IN"   OSMOSE.  21 

effecting  circulation  in  living  organisms,  but,  examined  criti- 
cally, what  essential  principle  does  either  represent  ? 

As  a  matter  of  fact,  they  express  nothing  further  than  that 
fluids  pass  through  membranes  and  traverse  interstices,  while 
they  leave  out  altogether  the  potential  factor  for  compelling 
them  to  do  so,  and  why  heat  should  energize  these  actions 
also  ;  not  to  mention  the  manner  of  connecting  them  with  the 
great  circulation  taking  place  in  the  globe  itself,  upon  which 
they  are  dependent  and  with  which  organic  unity  is  main- 
tained. At  the  same  time,  however,  we  have  to  note  the  curi- 
ous circumstance  that  physiologists,  in  seeking  to  explain 
osmotic  action,  have  constant  recourse  to  the  nomenclature  in 
polarity,  which  at  once  concedes  the  point  at  issue,  if,  for- 
sooth, language  means  anything  at  all.  In  fine,  the  endos- 
mometer  is  based  upon  the  '* affinity"  of  the  fluids  for  one 
another,  and  the  powers  of  "attraction"  exerted  on  the  part 
of  the  membranes  for  the  fluids.  And  should  ' '  affinity ' '  and 
'•attraction"  be  eliminated,  what  would  become  of  osmotic 
action  and  the  endosmometer  ?  Then,  again,  in  speaking  of 
diffusion,  which  is  essential  to  osmose,  we  note  language  of 
similar  import,  to  wit :  ' '  When  the  water  reaches  the  inner 
surface  of  the  membrane  (containing  alcohol),  it  instantly 
diffuses  itself  into  that  fluid,  partly  in  consequence  of  the 
mutual  repulsive  force  of  its  own  particles,  and  partly  from 
their  affinity  to  those  of  the  alcohol."  Italics  are  added. 
Further  comment  is  unnecessary. 

Now,  then,  in  regard  to  capillarity.  Is  it  not  true  that  water, 
after  passing  into  a  corner  of  a  dried  sponge  and  diffusing  it- 
self through  the  mass,  passes  out  again  at  the  surface  to  join 
the  great  mechanics  in  the  atmosphere,  expanding  and  rising 
up  in  this  by  what  is  known  as  evaporation  ?  while  heat  ener- 
gizes the  movement.  That  so  long  as  this  relation  of  the  parts 
shall  continue,  so  long  will  this  miniature  circulation  be  main- 
tained. Ceasing,  by  removal  of  the  sponge  from  the  water,  the 
former  speedily  becomes  dry  and  crisp  again,  as  the  flora  in 
iatal  drought. 

But  the  rapidity  with  which  water  at  first  enters  the  sponge  ! 
— how  is  that  circumstance  to  be  explained  %  The  answer  is  also 
readily  given  :     The  old  relationship  subsisting  between  them 


22  POLAR  ACTION   IN   CAPILLARITY. 

being  broken  up,  asserts  itself  the  first  opportunity  that  pre- 
sents, the  water  rushing  into  the  sponge  under  the  powerful 
action  of  the  polar  forces  intensified  by  electrical  induction, 
the  atmosphere  being  more  highly  electrical ;  otherwise  it  would 
not  be  a  gas.     The  dried  sponge,  then,  compels  water  into  it- 
self with  great  avidity  when  connection  is  made,  not  by  reason 
of  the  canal  system  and  interstices  existing  in  it,  which  facili- 
tate the  transit  only,  but  because  of  the  energy  in  the  polar 
forces  of  its  molecules,  which  are  highly  charged  with  oppo- 
site electricity,  the  aqueous  molecules  passing  on  immediately 
the  affinities  are  satisfied,  to  make  room  for  those  pressing 
on  behind  in  ceaseless  procession,  passing  out  at  the  surface 
again  into  the  great  mechanics  ;  the  supply  ceasing,  the  sponge 
speedily  becomes  dry  and  crisp  again,  the  pulling '-force  in  the 
atmosphere  and  the  pushing-force  in  the  sponge  produced  by 
repulsion,  compelling  it  all  out  to  the  very  last  molecule     it  is 
all  simple  enough.     In  fine,  all  these  actions  must  be  accounted 
for  in  a  theory  of  circulation,  and  if  polar  force  were  ex- 
cluded, it  would  be  utterly  impossible  to  explain  them.     Last, 
but  not  least,  with  the  extinction  of  the  solar  beam  as  the 
source  of  electrical  supply,  there  could  be  no  molecular  actions 
as  involved  in  the  movements  in  fluids,  consequently  there 
could  be  no  life,  showing  conclusively  that  the  principle  under- 
lying it  all  is  electrical  and  polar. 


CHAPTER  II. 

THE  FLORAL   CIRCULATION. 

Principle  in  the  Floral  Circulation— Action  of  the  Polar  Forces  and  the  Mode  of  Increas- 
ing it — Physiological  Experiment  Demonstrating  the  Energy  of  the  Polar  Forces  in 
Flora — Circulation  in  Flora  Comparatively  Slow— Deduction  to  be  Drawn  There- 
from—Explanation for  the  Rapidity  of  Circulation  in  Fauna  and  the  Amount  of 
Food  Consumed  by  Them. 

Perceiving  an  organic  connection  subsisting  between  circu- 
lation in  air  and  water  and  circulation  in  living  organisms,  we 
have  only  to  apply  the  preceding  principles  to  circulation  in 
trees  in  order  to  make  this  also  intelligible  and  easily  under- 
stood. 

For  example,  one  end  of  the  growth  is  rooted  in  the  earth, 
while  the  other  extends  to  variable  distances  in  the  atmos- 
phere, and  the  former  being  negative  and  the  latter  positive, 
the  conditions  obtain  for  producing  circulation  between  the 
two  ends.  Furthermore,  the  laws  of  electrical  induction 
should  tend  to  increase  this  action,  for  the  reason  that  the 
atmosphere  being  positive,  this  should  compel  the  positive 
electricity  in  the  bole  and  branches  into  'the  roots  and  root- 
lets and  pull  all  the  negative  electricity  in  the  latter  into 
the  bole  and  branches  and  terminal  leaflets  ;  and  being  thus 
differentiated  in  electrical  force,  the  highest  conditions  obtain 
for  producing  circulation  in-and-out  the  two  ends,  since  it 
would  effect  opposite  polarities  in  the  branches  and  the 
atmosphere  and  in  the  rootlets  and  the  juices  in  the  earth, 
thus  connecting  the  mechanics  through  and  through,  and 
producing  continuity  in  force,  which  the  scheme  calls  for. 
The  fluids  being  in  immediate  contact  with  the  cell- walls  in 
the  two  ends,  the  molecules  are  readily  drawn  through  the  thin 
places  in  the  walls  left  for  the  purpose,  and  being  once  in  "he 


24  POLAR   ACTION. 

channels  of  the  circulation,  pass  readily  up  and  down  the 
structure  under  the  action  of  the  polar  forces,  upon  the  same 
principle  precisely  as  obtains  for  producing  circulation  of  air 
and  wa  ter  in  the  globe  itself,  out  of  which  the  growth  is  evolved. 
Furthermore,  it  would  explain  the  enormous  force  which  is  in- 
volved for  producing  circulation  in  mammoth  trees;  e.  g.,  the 
giants  of  California,  some  of  which  are  said  to  be  four  hun- 
dred and  fifty  feet  in  height  and  ninety  feet  in  circum- 
ference, and  in  which  tons  upon  tons  of  sap  are  kept  in  motion 
from  end  to  end  of  the  mighty  growths,  since  the  force  which 
maintains  air  and  water  in  a  ceaseless  circuit  round  the  globe 
itself  is  commensurate  with  this  circumstance. 

Then,  again,  it  would  explain  the  oscillations  in  this  circu- 
lation with  the  waxing  and  waning  of  solar  force,  heat  being 
necessary  for  energizing  molecular  action  by  increasing  the 
polar  forces.  Thus,  with  the  return  of  spring,  the  vernal  heat 
and  moisture,  the  sap  leaps  up  the  tree,  and  the  whole  noise- 
less clockwork  is  again  set  in  motion,  the  buds  swelling  and 
bursting  into  foliage,  while  the  deep  sleep  of  winter  is  brought 
to  an  end  ;  the  matter  obviously  depending  upon  temperature 
for  energizing  the  polar  forces.  But  take  the  action  in  the  cell 
itself  as  further  example.  Thus  when  the  cell  of  a  succulent 
plant  is  examined  under  the  microscope,  a  distinct  circulation 
is  seen  to  be  taking  place  within  it,  the  fluids  passing  first  up 
one  side,  across  the  upper  end,  then  down  the  opposite  side  of 
the  cell  in  a  continuous  circuit  under  the  action  of  the  polar 
forces ;  a  miniature,  in  short,  of  the  great  circulation  taking 
place  in  the  globe  itself,  out  of  which  it  had  been  evolved. 
And  looking  thence  to  the  circulation  taking  place  in  the  plant 
itself,  it  is  at  once  perceived  that  this  also  is  but  an  extension 
of  the  same  principle  of  motion,  the  fluids  in  the  cell-empire, 
as  in  the  case  of  the  individual  cell,  passing  first  up  one  part, 
then  down  another  part  of  the  growth,  and  so  round  and 
round  the  limits  in  the  plant  in  a  continuous  circuit  under  the 
action  of  the  polar  forces,  while  heat  serves  for  energizing  the 
movement,  the  same  as  in  air  and  water. 

This  circumstance  may  be  seen  to  great  advantage  in  a  maple 
orchard  in  the  early  spring,  when  the  sap  is  running  up: 
With  the  object  of  obtaining  this  sap,  holes  one-half  inch  in 


ENEKGY   OF  POLAR  ACTION.  25 

diameter  and  about  one  and  one-half  inches  in  depth,  so  as  to 
penetrate  the  newest  wood,  are  made  in  the  bole  of  the  tree 
by  means  of  an  auger.  A  large  tree,  or  one  approximating 
two  feet  in  diameter,  contains  about  24  such  holes,  with  elder 
spouts  for  conveying  the  water  into  a  receiving  vessel  at  the 
foot,  commonly  an  excavated  section  of  a  tree  improvised  for 
the  purpose,  and  remaining  from  year  to  year,  though,  of 
course,  fresh  holes  are  made  every  year.  The  water  trickles 
through  these  spouts  in  a  tiny  stream.  Now,  then,  the  point 
we  wish  to  make  is,  that  during  night-time,  when  temperature 
falls,  the  flow  nearly  ceases,  and  when  this  approaches  the 
freezing  point,  it  stops  altogether ;  while  during  the  day-time, 
when  temperature  rises,  it  sets  in  again,  increasing  more  and 
more  as  day  advances,  till  about  3  p.  m.,  when  the  maximum 
is  reached,  which  corresponds  with  the  maximum  of  tempera- 
ture. From  this  time  on  it  grows  less  and  less,  with  the  de- 
cline of  solar  force,  till  night  sets  in,  when  the  sudden  fall  in 
temperature  brings  it  nearly  to  a  standstill.  On  a  bright,  warm 
day  following  a  cold  night  the  flow  is  very  rapid,  a  very  strik- 
ing illustration,  indeed,  of  the  intimate  relations  and  direct 
dependence  of  the  floral  circulation  upon  temperature,  and 
which,  of  course,  must  act  by  increasing  polarity  in  the  plant 
and  fluids  ;  otherwise  is  inexplicable.  Finally,  a  ready  means 
obtains  for  measuring  the  energy  of  the  polar  forces  in  the 
plant,  by  repeating  the  physiological  experiment  adopted  by 
Hales.  For  example,  he  discovered  "that  a  vine,  cut  in  the 
bleeding  season,  will  push  its  sap  up  a  glass  tube  as  high  as 
21  feet  above  the  surface  of  the  stump."  Now,  then,  since 
there  are  no  leaves  to  draw  upon  the  sap  by  evaporation,  it 
follows  that  the  lifting-force  to  this  great  column  of  sap,  and 
which,  of  course,  would  include  the  air  standing  upon  it  in  the 
open  end,  must  be  due  to  the  action  of  the  polar  forces  simply, 
no  other  force  applying  for  producing  it.  Some  evaporation 
takes  place  in  the  open  end,  it  is  true,  but  the  circulation  is 
greatly  in  excess  of  this,  the  sap  rapidly  accumulating  till  a 
column  21  feet  in  height  is  reached,  where  the  point  of  equi- 
librium is  reached,  and  a  balance  is  struck  between  the  action 
of  gravitation  and  the  action  of  the  polar  forces,  the  two  being 
antagonistic ;  the  weight   in  this  column  of  fluid  being  the 


20  POLAR  FORCES  IX  FLORA. 

measure  of  the  energy  of  the  polar  forces  in  the  terminal  ends 
or  poles  in  the  vine,  otherwise  is  inexplicable. 

Of  course,  evaporation  from  the  surface  of  the  leaflets  in- 
volves corresponding  absorption  in  the  rootlets,  for  maintaining 
a  balance  in  the  circulation,  in  this  manner  increasing  circula- 
tion in  proportion  to  the  evaporation  taking  place;  but  it  would 
not  explain  why  the  tree  fills  up  with  sap  before  there  are  any 
leaflets  and  out  of  which  the  leaflets  themselves  are  formed  ; 
while  it  leaves  unexplained  the  principle  in  evaporation  for 
connecting  the  internal  with  the  external  mechanics,  which  the 
scheme  calls  for,  continuity  in  force  demanding  this.  And  not 
water  only,  but  the  oxygen  which  is  disengaged  in  the  nutri- 
tive processes  from  carbonic  acid,  and  is  passing  out  from  the 
surface  of  the  leaflets  in  a  constant  stream,  the  molecules 
promptly  taking  up  their  relative  positions  in  the  atmosphere; 
while  carbonic  acid  is  compelled  in  the  tree  by  the  same  means, 
n  fine,  that  there  must  be  some  universal  and  all-pervad- 
ing principle  in  nature  for  connecting  the  prodigious  clock- 
work through  and  through,  and  bearing  upon  every  piece  to 
the  very  molecules  and  atoms  themselves,  since  it  is  through 
them  the  masses,  living  or  non-living,  are  built  up  and  sustained, 
or  are  pulled  down  and  again  redistributed.  There  is  no  diffi- 
culty, then,  in  accounting  for  circulation  in  flora  any  more  than 
there  is  in  accounting  for  circulation  in  the  air  and  water  out  of 
which  they  are  evolved,  while  all  the  phenomena  fall  readily 
into  line  in  regular  order  and  succession,  and  leave  no  out- 
standing quantity  refusing  absorption.  And  in  this  connec- 
tion we  should  not  fail  to  notice  a  deeply  suggestive  fact, 
namely  :  why  water  should  pass  out  at  the  leaflets  in  a  con- 
stant stream  by  evaporation,  and  if  not  as  rapidly  renewed  by 
a  similar  stream  passing  in  at  the  rootlets  for  maintaining  a 
balance,  the  plant  would  speedily  become  dry  and  crisp,  as  oc- 
curs in  fatal  droughts.  This  also  is  easily  answered.  Since 
water  is  the  medium  of  transportation  for  the  nutritive  and 
force-producing  elements,  it  follows  when  these  are  withdrawn 
by  the  nutritive  and  functional  processes,  that  a  correspond- 
ing proportion  of  water  should  be  discharged  in  order  to  make 
room  for  the  fresh  water  and  supplies  coming  into  the  organ- 
ism; the  one  involving  the  other,  as  in  no  other  way  could  a 


CIECDLATIOlSr   AND   NUTRITION.  27 

balance  bo  maintained,  water  being  the  medium  of  transporta- 
tion Otherwise  nutrition  and  function  would  have  arrest 
from  the  absence  of  the  gases  and  the  alkaline  earths  and 
minerals  dissolved  in  it  by  action  of  the  polar  forces.  In  the 
fauna  a  comparatively  large  amount  of  water  is  being  thus 
discharged  from  the  surface  for  the  purpose  of  regulating 
body- temperature,  for  which  a  physiological  balance  obtains 
in  the  organism,  being  the  most  effective  means  of  doing  this, 
a  high  temperature  being  inimical  to  life.  Thus  during  exer- 
cise, which  tends  to  put  up  temperature,  the  skin  becomes 
flushed  for  cooling  the  blood;  at  the  same  time  is  bathed  in  per- 
spiration. But  this  matter  is  more  fully  treated  in  another  place. 

Concerning  the  Rapidity  of  the  Floral  Circulation  and  the 
Deduction  to  be  drawn  therefrom :  The  fluids  having  to  pass 
from  cell  to  cell  by  means  of  thin  places  in  the  walls  left  for 
the  purpose,  each  cell  forming  a  separate  and  distinct  com- 
partment in  itself,  it  follows  that  circulation  in  flora  is  com- 
paratively slow.  Still,  this  is  sufficient  for  the  nutritive 
processes  simply,  which  require  a  slow  circulation  to  afford 
opportunity,  so  to  speak  for  crystallizations  to  take  place  in 
the  formative  processes ;  whereas,  in  animal  life,  the  very 
opposite  obtains,  the  purpose  here  being  the  generation  of 
force,  which  is  expended  in  producing  the  multitudinous  ac- 
tions taking  place  in  them,  since  everything  is  evolved  out  of 
circulation,  which  is  made  commensurate  with  the  special 
physiological  requirements.  Hence,  the  differentiation  of  a 
vascular  system,  together  with  the  special  arrangements  that 
obtain  in  them  for  compelling  circulation  to  be  in  correspond- 
ence with  the  amount  of  force  which  is  expended  in  producing 
the  various  movements,  and  which  are  classified  as  voluntary 
and  involuntary,  or  animal  and  organic,  but  all  of  which  are 
directly  dependent  upon  circulation,  as  must  appear  obvious. 

The  point  we  wish  to  make  is,  that  the  nutritive  processes 
require  a  slow  circulation,  and  in  proof  of  this  fact,  we  have 
overwhelming  evidence  furnished  in  the  flora,  notably  in  the 
case  of  a  pumpkin  weighing  265  lbs.,  which  was  produced  in 
a  period  of  less  than  60  days  from  the  time  of  flowering,  and 
that  upon  a  vine  but  a  little  more  than  an  inch  in  diameter.* 

*  All  the  buds  are  pinched  off  save  the  one  selected. 


28  CIRCULATION   AND   FORCE. 

In  succulent  plants,  such  as  grasses,  peas,  beans,  etc,  growth  is 
very  rapid.  And  in  many  others  as  well ;  e.  g. ,  melons,  goiu-ds, 
roots  or  tubers.  The  fruits  also.  So,  then,  nutrition  per  se  does 
not  require  a  rapid  circulation,  and  sufficient  only  to  effect  the 
accretions  and  crystallizations  for  producing  growth,  inclusive 
of  the  arrangements  in  the  structures  for  prod  ucing  motion  which 
animal  life  involves.  And  we  make  the  deduction  for  animal 
life,  for  the  reason  that  the  principle  in  nutrition  is  the  same, 
and  must  be  so  in  the  very  nature  of  things.  Namely, 
the  action  of  the  polar  forces  for  pulling  the  molecules 
into  position  and  locking  them  in  the  structures,  a  slow 
circulation  favoring  this  action.  Consequently,  nutrition  goes 
on  most  rapidly  during  sleep  and  repose,  when  circulation  is 
slowest.  But  for  evolving  force  in  the  organism  for  producing 
the  multitudinous  actions  taking  place  in  the  body,  calls  for 
a  rapid  circulation  to  make  it  commensurate  with  this  circum- 
stance, the  two  being  necessarily  in  correspondence,  for  in  no 
other  way  could  a  balance  be  maintained. 

It  calls  for  considerable  expenditure  of  force ;  hence,  the 
amount  of  food  consumed,  especially  by  warm-blooded  animals, 
the  maintenance  of  temperature  alone  calling  for  large  supplies, 
as  this  approximates  100  degrees  Fahr.;  in  many  animals, 
birds  especially,  it  runs  a  number  of  degrees  higher.  In  this, 
too,  there  is  variety,  as  in  everything  else,  animate  and  inani- 
mate, variety  being  the  one  pronounced  circumstance  in 
Nature. 

This  brings  us  to  the  means  for  increasing  the  animal 
circulation,  commensurate  with  the  force  which  is  expended  in 
them. 


CHAPTER   III. 

THE    ANIMAL    CIRCULATION. 

Principle  in  the  Animal  Circulation — Adjustment  with  Pressure  and  the  Power  of  Pro- 
ducing Rapid  Rhythmical  Changes  in  Pressure,  the  Law  in  the  Animal 
Circulation — The  Movements  in  Respiration  :  Heart,  Arteries,  etc.,  Pumping- 
Actions  for  Increasing  Circulation,  the  Whole  Forming  a  Connected  Movement  for 
Increasing  Circulation  Between  the  Cell-Brood  and  Environment,  whence  the  Nutri- 
tive and  Force-Producing  Elements  are  Derived,  and  into  which  the  Waste  Products 
are  Returned — Explanation  for  the  Correlation  of  the  Vaso-Motor  and  Voluntary- 
Motor  Centres  with  the  Respiratory  Centre — Mechanics  in  Inspiration  and  Expira- 
tion— Two  Respiratory  Movements  Performing  at  the  Same  Time  ;  One  in  the 
Lungs,  the  Other  in  the  Tissues — Physiological  Experiments,  Showing  that  Pressure 
is  the  Fundamental  Circumstance  in  the  Animal  Organism,  with  which  Everything 
has  Adjustment,  and  that  the  Actions  in  the  Lungs,  Heart,  Vessels  and  Hollow 
Viscera  Relate  to  Changes  in  Pressure  for  Compelling  Movement  in  the  Contents  : 
i.  e.,  for  Increasing  Circulation — Otherwise  are  Meaningless. 

We  are  now  in  a  position  to  take  up  the  animal  circulation 
and  explain  the  phenomena  appertaining  to  it ;  notably,  the 
actions  in  respiration  and  in  the  heart  and  blood  vessels, 
together  with  the  whole  scheme  in  animal  structure  and  func- 
tion. 

But  in  addition  to  the  mechanics  for  effecting  circulation 
in  plants — namely,  the  action  of  the  polar  forces — we  will  now 
have  to  introduce  the  principle  of  change  in  pressure,  before 
alluded  to  for  increasing  circulation  in  the  air,  only  that  the 
action  is  here  confined  within  narrow  limits  and  is  rapidly 
repeated,  said  circumstance  being  announced  by  the  pumping 
movements  in  respiration  and  in  the  heart  and  blood  vessels, 
which  relate  to  rhythmical  changes  in  pressure  for  increasing 
circulation  commensurate  with  the  physiological  requirements, 
hence  rising  and  falling  in  correspondence  with  these ;  other- 
wise are  meaningless.     In  short,  the  animal  circulation  is 

BASED  UPON  PRESSURE  AND  THE  POWER  OF  PRODUCING  RAPID 
RHYTHMICAL  EXPANSIONS  AND  CONTRACTIONS  IN  THE  ORGANS 
FOR  CHANGING   PRESSURE,    WHEREBY   THE   COMMERCE   IS   COM- 


30  THE  LAW  IN  THE  ANIMAL   CIRCULATION. 

PELLED  IN  THE  VESSELS  AND  THROUGH  THE  TISSUES  WITH  COR- 
RESPONDING ENERGY,  THE  FLUIDS  FLOWING  FROM  HIGH  TO 
LOW  PRESSURE   IN   CONFORMITY   WITH    ORGANIC    LAW.      Hence 

the  mechanical  principle  in  respiration  for  compelling  the  air 
in  and  out  of  the  lungs,  is  the  same  as  in  the  heart  and  blood 
vessels  for  speeding  the  blood  between  the  lungs  and  the  cell- 
brood,  the  workmen  in  the  tissues  and  objective  point  of  all 
the  supplies.  Moreover,  they  form  a  connected  movement, 
as  in  no  other  way  could  a  balance  be  maintained.  In  other 
words,  the  pumping  action  in  the  lungs  and  abdomen  for  com- 
pelling the  force-producing  and  nutritive  elements  in  the  vas- 
cular channels  calls  for  these  pumping  actions  in  the  heart 
and  vessels  for  circulating  them,  for  in  no  other  way  could  it 
be  produced.  And  but  for  this  power  of  producing  rapid 
rhythmical  changes  in  pressure  in  the  organs  and  tissues  for 
increasing  circulation  development  would  inevitably  have  arrest 
at  the  iiora,  since  no  other  means  obtains  for  increasing  circu- 
lation commensurate  with  the  physiological  requirements  in 
animal  organisms  ;  while  the  speed  of  the  currents  thus  pro- 
duced has  determination,  by  the  rapidity  and  energy  of  the 
pumping  actions  taking  place  in  the  organs,  and  pervading  the 
entire  body,  exclusive  of  the  bones  and  cartilages  (the  frame- 
work of  support  to  the  soft  tissues),  in  which  circulation  is 
analogous  with  that  in  plants.  And  while  the  scheme  is 
comprehensive,  involving  multitudinous  arrangements  in  the 
organs  and  tissues  for  producing  the  actions  and  for  unifying 
them,  still  it  is  all  simple  enough  and  easily  understood.  Of 
course,  it  would  naturally  include  the  digestive  and  assimi- 
lative functions,  with  the  secretory  and  excretory,  for  it  all 
relates  to  circulation  and  the  maintenance  of  a  balance  in 
the  organism.  And  since  pressure,  by  reason  of  the  com- 
pressibility of  the  tissues,  is  transmitted  through  the  body 
upon  all  the  organs  to  the  minutest  histological  elements  and 
cells,  it  is  at  once  made  available  for  increasing  circulation, 
commensurate  with  the  physiological  requirements.  Hence, 
the  universal  pumping  actions  taking  place  in  the  body,  as  be- 
fore remarked.  This  necessity  for  increasing  circulation  in  the 
fauna,  over  and  above  what  takes  place  in  the  flora,  in  order  to 
make  it  commensurate  with  the  force  which  is  expended  in 


THE   LAW   IN   THE   ANIMAL   CTECTJLATION.  31 

them,  is  sufficiently  obvious  ;  since  every  variety  of  motion 
involves  a  corresponding  expenditure  of  force  for  producing 
it,  while  this  in  turn  is  evolved  from  the  chemical  combina- 
tions of  substances  which  are  carried  into  the  organism  by 
circulation — notably  oxygen  and  carbon — the  waste  products 
being  carried  out  by  the  same  means.  Hence,  the  greater  the 
expenditures  of  force,  or  the  more  rapid  the  voluntary  move- 
ments, the  more  rapid  are  the  actions  in  the  lungs,  heart  and 
blood-vessels  for  making  circulation  commensurate  with  this 
circumstance,  the  whole  being  in  correspondence,  for  in  no 
other  way  could  a  balance  be  maintained.  Thus,  in  the  recum- 
bent position,  the  action  in  the  lungs,  heart  and  blood-vessels 
is  lowest ;  but  the  instant  the  erect  position  is  reached,  there  is 
considerable  increase  in  the  action  for  promptly  supplying  the  in- 
creased expenditures  which  this  calls  for,  while  there  is  progress- 
ive increase  in  correspondence  with  the  swell  in  the  activities, 
therefore  is  greatest  when  the  animal  is  running.  In  other 
words,  pressure  is  invoked  in  the  measure  of  the  requirements 
by  means  of  these  pumping  actions.  But  in  the  absence  of 
this  law  of  pressure  upon  which  to  base  the  mechanics,  it  were 
as  utterly  impossible  to  explain  the  relative  phenomena,  ana- 
tomical and  physiological,  as  it  would  be  to  account  for  the 
strong,  bony  skeleton,  with  the  powerful  levers  and  muscles, 
for  effecting  locomotion  in  land  animals  in  the  absence  of 
gravitation  for  compelling  this  circumstance ;  the  one  no  more 
than  the  other.  Proceeding  upon  this  line  of  investigation, 
then,  we  shall  pass  in  brief  review  the  scheme  in  the  animal 
circulation,  thence  to  the  special  phenomena  in  detail,  in 
which  will  be  clearly  shown  the  admirable  simplicity  and  mar- 
velous perfection  which  characterize  the  mechanics  through- 
out, and  why  everything  is  just  as  it  should  be  for  producing 
the  mechanical  work  involved  in  the  circulation,  thus  furnish- 
ing overwhelming  proof  of  the  correctness  of  the  premises, 
making  the  argument  unanswerable.  We  begin  with  the 
nervous  centres  for  operating  the  mechanics  and  unifying  the 
movements,  in  itself  furnishing  the  highest  order  of  evidence 
in  support  of  the  theory,  being  means  to  ends. 


32  CORKELATION  OF  NERVOUS  CENTRES. 

Fjjplanation  for  the  Existence  and  Correlation  of  the  Three 
Great  Nervous  Centres  at  the  Base  of  the  Brain  in  the 
Medulla  Oblongata,  notably  Respiratory,  Vaso-Motor 
and  Voluntary-Motor  Centres: 

This  relation  which  the  animal  organism  sustains  to  pressure 
enables  us  to  offer  a  scientific  explanation  for  the  existence 
and  correlation  of  the  three  great  nervous  centres  at  the  base 
of  the  brain  in  the  medulla  oblongata  for  operating  the 
mechanics,  or  the  respiratory,  vaso-motor  and  voluntary-motor 
centres  ;  otherwise  inexplicable,  and  which  is  briefly  as  follows: 

Respiratory  Centre. — Since  respiration  is  the  great  pump- 
ing action  for  compelling  the  fluids  in  the  organism,  it  is 
manifest  that  this  calls  for  a  common  nervous  centre  for  pro- 
ducing and  coordinating  the  action  in  the  lungs  and  con- 
taining walls,  to  the  end  that  air  and  blood  may  be  pumped 
through  the  lungs  for  respiratory  purposes,  and  which  would 
also  include  the  force-producing  elements  in  the  intestinal 
canal,  since  it  is  by  chemical  combination  of  oxygen  with 
carbon  that  force  is  generated,  a  measure  of  oxygen  calling 
for  a  measure  of  carbon  in  order  to  effect  it.  Hence,  this 
respiratory  centre,  together  with  the  large  pneumogastric 
nerves  extending  thence  into  the  abdomen  for  connecting  and 
coordinating  the  viscera  with  respiration.  Of  course,  there 
must  be  a  philosophic  reason  for  these  nervous  combinations, 
or  one  based  upon  organic  law,  as  they  are  persistent  and  uni- 
versal. It  does  not  follow  that  heat  is  generated  in  the  lungs, 
which  is  not  the  case,  only  that  intestinal  absorption  should 
be  in  correspondence  with  respiration  for  maintaining  a  balance, 
otherwise  impossible. 

Vaso-Motor  Centre. — Several  reasons  make  the  existence  of 
the  vaso-motor  centre  an  absolute  necessity,  namely:  1.  It  is 
necessary  for  coordinating  the  blood-vascular  system  with 
respiration  in  order  to  produce  an  uninterrupted  current  of 
the  blood  from  the  lnngs  to  the  tissue  territories  for  maintain- 
ing a  balance  between  supply  and  demand  ;  otherwise  this 
could  not  be  done.  In  other  words,  the  pumping  actions  in 
the  heart  and  blood  vessels  must  have  adjustment  with  the 
pumping  actions  in  the  lungs  for  producing  correspondence, 


CORRELATION   OF   NERVOUS   CENTRES.  33 

the  one  necessarily  having  adjustment  with  the  other,  as  must 
appear  obvious.  Hence,  the  existence  of  this  centre,  together 
with  its  correlation  with  the  respiratory,  lor  in  no  other  way 
could  continuity  in  force  be  produced.  2.  Since  the  vessels 
expand  and  contract  upon  their  contents  for  increasing  or 
diminishing  the  blood  for  regulating  the  local  actions  which 
are  ever  changing,  it  follows  that  there  must  be  a  means  for 
limiting  the  local  supplies  and  regulating  the  movements 
throughout  so  as  to  maintain  a  balance ;  otherwise,  some 
parts  of  the  system  would  have  more  blood  than  necessary, 
while  other  portions  would  suffer  from  dangerous  ansemia ; 
hence,  this  vaso-motor  centre  for  compelling  circulation  in  the 
measure  of  the  requirements.  3.  Last,  but  not  least,  the  hign 
pressure  in  the  arterial  system  for  increasing  circulation  in 
the  capillaries  tends  to  produce  dangerous  accumulations  in 
the  venous  system,  where  pressure  is  low,  and  since  there  is  a 
norm  of  blood  which  is  kept  in  constant  motion,  it  follows 
there  must  be  some  means  for  regulating  the  capacity  of  the 
venous  system,  which  is  something  like  four  times  as  great  as 
the  arterial ;  hence  this  vaso-motor  centre  for  regulating  that 
circumstance  also.  Finally,  the  whole  is  set  to  respiration, 
for  this  is  the  great  pendulum  movement  in  the  clockwork 
with  which  everything  must  have  adjustment,  since  it  is  the 
means  for  compelling  the  commerce  in  the  organism,  while  the 
heart  and  vessels  function  as  a  carrier. 

3.  Voluntary-Motor  Centre. — The  existence  of  the  vol- 
untary-motor centre,  together  with  its  correlation  with  the 
respiratory  and  vaso-motor  centres,  is  also  easily  understood. 
Thus,  since  every  movement  involves  a  corresponding  expendi- 
ture of  force  for  producing  it,  and  this  is  evolved  out  of  circu- 
lation from  substances  brought  into  the  organism  by  this 
means,  it  follows  that  respiration  and  the  actions  in  the  heart 
and  vessels  must  be  in  correspondence  with  the  voluntary 
movements  in  order  to  maintain  a  balance  between  supply  and 
demand,  for  in  no  other  way  could  this  be  done ;  hence  the 
correlation  of  this  nervous  centre  with  the  other  two  centres 
so  as  to  produce  continuity  in  force,  which  the  scheme  calls  for. 

In  fine,  respiration  and  circulation  together  form  the  basis 
of  the  activities,  consequently  the  latter  must  have  adjust- 


34  CORRELATION   OF   NERVOUS   CENTRES. 

ment  with  the  former.     Hence  the  correlation  of  these  nervous 
centres  in  the  medulla  oblongata. 

The  necessity  for  producing  and  coordinating  the  voluntary 
movements  would,  of  course,  explain  the  existence  of  the 
voluntary-motor  centre,  but  at  the  same  time  it  would  not 
account  for  the  correlation  of  this  centre  with  the  other  tw<  > 
centres.  In  this  manner,  then,  that  vexed  problem  in  physi- 
ology has  scientific  explanation,  and  it  is  at  once  seen  these 
nervous  combinations  in  the  medulla  oblongata  are  inevitable 
from  the  very  nature  of  things,  the  law  in  the  circulation  and 
continuity  in  force  alike  compelling  this  circumstance. 

Thus  with  a  common  law  underlying  it  all,  and  the  same 
principle  in  mechanics  for  increasing  circulation — i.  e.,  rhyth- 
mical changes  in  pressure — it  is  manifest  that  correspondence 
in  the  structures  is  also  made  inevitable,  or  that  similar 
arrangements  should  obtain  in  the  animals  for  producing  cir 
dilation  and  the  voluntary  movements.  And  this  would  include 
not  only  the  systemic  mechanics,  but  the  anatomical  disposi- 
tions in  the  walls  of  the  organ  as  well ;  while  this  again  would 
have  determination  by  the  character  of  the  work,  the  nature  of 
the  contents,  and  the  amount  of  force  which  is  required  for  com- 
pelling movement  in  them  in  the  rule  of  the  special  functions, 
so  that  all  is  readily  explained  and  made  intelligible.  Indeed, 
this  principle  for  increasing  circulation  must  apply  to  the 
very  cells  themselves,  which  expand  and  contract  under 
stimulus  for  producing  afflux  and  efflux  of  the  fluids  in  which 
they  are  submerged,  for  in  no  other  way  could  circulation  be 
increased  within  the  cells  for  increasing  metabolism. 

Finally,  each  organ  by  possessing  separate  and  independent 
local  nervous  centres  appertaining  to  its  own  special  functions, 
but  connecting  with  the  systemic  apparatus,  enables  the  local 
actions  to  be  increased  or  diminished  as  occasion  may  require, 
•without  interfering  with  the  general  functions.  As  the  me- 
chanics are  fully  brought  out  and  explained  at  the  proper 
time  and  place  in  the  text,  it  will  not  be  necessary  at  this 
early  stage  to  do  more  than  briefly  refer  to  the  cardinal  cir- 
cumstances in  order  to  prepare  the  mind  of  the  student  for  the 
radical  changes  introduced  in  present  physiology,  and  which 
lie  will  at  once  see  are  not  only  logical  and  necessary,  but  un- 


MECHANICAL   PRINCIPLE  IN   RESPIRATION.  35 

avoidable.  First  of  all,  however,  let  Mm  get  himself  right  by 
emancipating  and  disenthralling  his  mind  of  all  prejudice 
tending  to  mar  the  judgment  and  obscure  the  mental  vision,  to 
the  end  that  he  may  listen  to  reason  and  weigh  evidence  dis- 
passionately; otherwise,  at  very  best  he  is  but  partly  rational, 
his  opinion  to  be  little  depended  upon  and  unworthy  of  respect. 
After  he  has  done  that,  let  him  lay  firm  hold  of  this  law 
of  pressure  underlying  the  animal  organism  if  he  would  under- 
stand the  phenomena  in  animal  mechanics,  since  they  all  relate 
to  this  fundamental  force  in  nature.  Now,  then,  it  being  true 
the  body  has  special  adjustments  with  pressure,  while  the  law 
in  the  circulation  consists  in  rapid  rhythmical  changes  in 
pressure,  it  remains  to  inspect  the  phenomena  from  this  stand- 
point, commencing  with  respiration,  the  initial  action  for  com- 
pelling the  commerce  in  the  organism,  as  before  remarked. 

Import  of  Inspiration  and  Expiration. — Briefly,  Inspira- 
tion is  the  effort  to  develop  a  lower  pressure  in  the  lungs 
than  exists  externally ;  in  consequence,  the  air  and  blood 
flow  into  the  low-pressure  area  till  pressure  is  uniform,  the 
amount  so  flowing  in  being  the  exact  measure  of  lung  ex- 
pansion ;  no  more  and  no  less :  while  Expiration  is  the 
effort  to  develop  a  higher  pressure  in  the  lungs  than  exists 
externnlly,  when  the  air  and  blood  in  consequence  flow  out  of 
the  organs  till  pressure  is  again  uniform,  the  amount  so  flow- 
ing out  being  the  exact  measure  of  lung  contraction  ;  no  more 
and  no  less.  In  this  manner,  then,  a  dual  circulation  of  air 
and  blood  is  maintained  through  the  lungs  for  respiratory 
purposes,  the  former  flowing  in  by  way  of  the  trachea  and 
the  latter  through  the  right  side  of  the  heart  and  pulmo- 
nary artery  during  inspiration  ;  but  during  expiration  the  air 
flows  out  by  reflux  action  through  the  route  of  ingress  to 
the  environment,  while  the  blood  itself  passes  through  the 
four  pulmonary  veins  into  the  left  side  of  the  heart  and 
arterial  system  on  its  way  to  the  cell-brood  in  the  tissues. 
But  the  great  inertia  in  the  blood  calls  for  the  expenditure  of 
considerable  force  for  bringing  it  into  correspondence  with  the 
circulation  of  air  in  the  alveoli,  the  obvious  purpose  being 
to  maintain  fresh  air  and  venous  blood  in  close  proximity, 
for    effecting    mutual    interchange.       Hence    the    numerous 


36  MECHANICAL   PRINCIPLE   IN    RESPIRATION. 

muscles  and  nerves  in  the  heart  and  blood  vessels  for  increas- 
ing circulation  in  the  lungs  and  in  the  tissues  and  for  coordinat- 
ing them  with  the  actions  taking  place  in  both  localities,  a  cir- 
cumstance which  is  fully  brought  out  further  on.  But  in  order 
to  produce  these  dual  circulations  of  air  and  blood  through  the 
alveoli,  it  is  absolutely  necessary  for  the  lungs  and  containing 
walls  to  expand  and  contract  together  or  simultaneously,  for 
in  no  other  way  could  the  requisite  changes  in  pressure  be 
produced  within  the  alveoli  for  compelling  afflux  and  efflux  of 
these  fluids,  neither  could  a  balance  in  pressure  be  maintained 
in  the  pleurse  for  obviating  effusions  in  these  cavities,  other- 
wise inevitable  ;  and  which  should  speedily  put  an  end  to 
respiration  by  preventing  lung  expansion  ;  such  concert  of 
action  in  the  parts  being  effected  by  the  nervous  apparatus, 
notably  the  pneumogastric,  intercostal  and  phrenic  nerves 
and  the  so-called  organic  or  sympathetic  system,  which  is 
intimately  connected  with  the  lungs,  and  undoubtedly  serving 
for  coordinating  these  actions.  The  mechanics  is  compre- 
hensive but  easily  understood,  the  law  in  the  circulation  com- 
pelling these  arrangements  for  maintaining  a  balance  and 
keeping  the  fluids  within  their  channels  ;  otherwise  impossible, 
since  they  flow  from  high  to  low  pressure  in  conformity  with 
universal  law.  And  all  these  nerves  being  correlated  in  the 
medulla  oblongata,  harmonious  action  throughout  is  readily 
produced.  And  which,  of  course,  would  include  the  vaso- 
motor system,  together  with  the  nerves  to  the  heart  for 
bringing  the  blood  into  correspondence  with  the  circulation  of 
air  in  the  lungs.  In  this  manner,  then,  these  wonderful  nervous 
ccmbinations  in  the  medulla  oblongata  are  readily  explained, 
and  they  can  be  exiuained  in  no  other  way.  And  not  this 
circumstance  only,  but  all  the  anatomical  dispositions  which 
obtain  in  the  organs  for  producing  these  actions  as  well,  being 
means  to  ends  simply  ;  furthermore,  they  cannot  be  explained 
in  any  other  way.  Thus  everything  is  in  correspondence,  and 
nothing  is  left  out.  And  this  should  be  the  case,  since  the 
mechanism  is  necessarily  founded  upon  law ;  hence  every  piece 
in  the  comprehensive  clockwork  must  have  definite  adjust- 
ments and  relations  with  the  rest,  as  must  appear  obvious. 
In  short,  a  physiology  which  does  not  explain  anatomy  is  no 


LUNGS   AND   VENOUS    SYSTEM.  37 

physiology  at  all.  But  in  the  absence  of  the  fundamental 
principle  upon  which  the  mechanics  is  based,  of  course  this 
would  be  impossible. 

The  animal  organism  being  founded  upon  pressure,  while 
the  law  in  the  circulation  consists  in  rapid,  rhythmical 
changes  in  pressure,  as  alleged,  it  follows  that  the  pumping 
action  in  the  lungs  should  compel  simultaneous  afflux  and 
efflux  of  air  and  blood  in  the  alveoli,  since  both  fluids  freely 
communicate  with  these  chambers  through  a  special  tubular 
system  arranged  for  the  purpose,  the  heart  not  interfering 
with  but  rather  assisting  this  mechanics  by  putting  additional 
force  upon  the  blood,  and  which  its  greater  inertia  calls  for,  as 
before  remarked ;  at  the  same  time  the  heart  itself  is  coordi- 
nated with  the  lungs,  both  doing  inspiration  and  expiration, 
so  that  in  nowise  is  the  action  in  the  lungs  interfered  with  by 
the  action  in  the  heart,  the  special  functions  in  which  will  be 
described  later  on,  and  their  true  relation  to  the  circulation 
fully  shown. 

The  following  diagram  of  the  venous  system,  showing  the 
relations  it  sustains  to  the  lungs  (Fig.  7),  will  serve  for  im- 
pressing the  matter  For  examine,  the  air  {L)  and  the  venous 
blood  (A,  A,  A)  freely  communicate  with  the  alveoli ;  hence, 
when  the  lungs  expand  during  inspiration  for  reducing  press- 
ure in  the  alveoli,  the  air  and  blood,  coming  from  opposite 
directions,  must  flow  into  these  chambers  simultaneously  till 
pressure  is  uniform,  pressure  compelling  them  to  do  so.  Upon 
the  other  hand,  when  the  lungs  contract  during  expiration 
for  increasing  pressure  in  the  alveoli,  the  air  and  blood 
must  flow  out  of  these  compartments  simultaneously  till 
pressure  is  again  uniform,  the  one  by  reflux  action  through 
the  route  of  ingress,  the  other  into  the  left  chambers  of  the 
heart  and  arterial  system,  the  valves  at  the  right  side  of 
the  heart  compelling  this  circumstance  by  preventing  reflux, 
as  in  the  case  of  the  air  in  the  air  passages.  From  this 
arrangement,  then,  it  is  very  readily  perceived  that  currents 
of  air  and  blood  should  flow  in  and  out  of  the  alveoli 
simultaneously  during  respiration,  the  one  necessarily  involv- 
ing the  other.  But  to  this  we  must  add  the  arrangements 
in  the  heart  and  vascular  system  for  bringing  the  blood  into 


38 


LUNGS   AND  VENOUS   SYSTEM. 


correspondence  with  the  circulation  of  air  in  the  alveoli,  there 
being  a  given  measure  of  each,  for  which  special  adjustments 
obtain  in  the  compartments,  as  also  for  producing  an  uninter- 
rupted flow  of  blood  through  the  lungs,  and  the  whole  scheme 
in  respiration  will  be  readily  apprehended.  First  of  all,  let 
it  be  understood  that  the  generation  of  force  in  the  organism 


j 


Fig.  7. — Diagrammatic  representation  of  the  Venous  System,  showing  the  relations  it 
sustains  to  the  Lungs.  A,  A,  A,  venous  system  represented  as  pyramids,  with  the 
bases  in  the  tissues  and  the  apices  at  the  heart  and  lungs  (E,  C)  ;  B,  pulmonary- 
artery  ;  D,  trachea  ;  F,  diaphragm  ;  L,  atmosphere. 

is  the  purpose  of  all  these  arrangements,  and  this  can  only  be 
done  by  means  of  the  pumping  actions  in  the  lungs,  with 
which  everything  must  have  adjustment ;  hence  would  include 
the  intestines  with  the  blood-vascular  and  lymphatic  systems, 


TWO   EESPIRATORY   MOVEMENTS.  39 

since  it  all  relates  to  circulation  and  the  supply  of  the  cell- 
brood  in  the  tissues,  through  whose  agencies  the  special 
phenomena  are  evolved. 

Finally,  we  have  to  mention  the  existence  of  two  respiratory 
movements  performing  at  the  same  time  in  the  body,  notably 
one  in  the  lungs,  the  other  in  the  tissues  ;  the  former  for  pump- 
ing the  commerce  in  the  organism,  the  latter  for  pumping  it  in 
and  out  of  the  tissues  for  the  due  supply  of  the  cell-brood  and 
removal  of  waste  products  ;  while  the  vascular  system,  inclu- 
sive of  the  heart,  functions  as  a  carrier  between  these  two 
poles  in  the  circulation,  with  which  they  are  coordinated  by 
means  of  the  nerves  extending  over  them  and  connecting  them 
with  the  medulla  oblongata,  the  vaso-motor  and  respiratory 
centres,  whence  force  is  propagated  over  all  the  structures,  to 
the  end  that  a  current  of  blood  may  be  maintained  between 
the  cell-brood  and  environment  for  the  due  supply  of  the 
nutritive  and  force-producing  elements  and  removal  of  waste 
products.  The  composite  character  in  the  arterial-  tracings,  or 
the  existence  of  respiratory,  cardio-arterial  and  dicrotic  or 
capillary  waves,  which  are  superposed,  one  upon  the  other,  in 
the  order  named;  or  the  respiratory,  by  the  cardio-arterial,  and 
the  latter  by  the  capillary  or  dicrotic  waves,  is  to  be  explained 
by  this  circumstance.  In  fine,  these  waves  or  respiratory  pul- 
sations are  throbbed  over  the  vessels  from  the  respiratory 
centre  in  the  medulla  oblongata,  by  means  of  the  nervous 
plexuses,  extending  over  the  vessels.  "Traube's  Curves," 
together  with  the  physiological  problem  connected  with  the 
curves  in  arterial  and  intra-thoracic  pressure,  are  readily 
explained.  By  means  of  this  law  of  pressure  underlying  the 
organism,  then,  all  this  is  easily  accounted  for  and  made  intelli- 
gible; otherwise  is  utterly  inexplicable.  The  most  incontro- 
vertible evidence  of  rhythmical  expansions  and  contractions 
taking  place  in  the  arteries  and  capillaries,  synchronous  with 
respiration  and  the  action  in  the  heart,  is  submitted,  placing 
this  function  in  the  vessels,  beyond  the  shadow  of  a  doubt  or 
the  possibility  of  mistake,  at  the  same  time  the  common  rela- 
tion which  it  all  sustains  to  pressure  is  too  obvious  for  con- 
troversy. 

Last  but  not  least,  we  have  to  note  the  deeply  significant  fact 


40  ARTERIAL   PRESSURE  AND   DEVELOPMENT. 

of  an  increase  of  pressure  in  the  arterial  system  with  pro- 
gress in  development,  notably  in  warm-blooded  animals.  Thus, 
when  a  cold-blooded  animal  is  decapitated,  the  blood  issues 
very  slowly  out  of  the  open  ends  of  the  vessels,  oozes  and 
wells  out  simply,  trickling  down  in  a  tiny  stream  upon  the 
ground,  dark  and  venous  in  appearance  ;  whereas  in  the  case 
of  a  warm-blooded  animal  it  spurts  out  to  a  considerable  dis- 
tance from  the  body  in  a  sustained  stream,  which  issues  per 
saltam  or  in  a  succession  of  leaps,  the  latter  being  due  to 
rhythmical  contraction  in  the  vessels  for  increasing  pressure, 
while  the  former  is  due  to  the  permanent  high  pressure  which 
exists  in  the  arterial  system  for  increasing  circulation  in  the 
tissues,  at  the  same  time  it  functions  as  the  ms  a  tergo  to  the 
venous  system  for  compelling  the  venous  blood  toward  the 
lungs,  thereby  assisting  the  action  in  those  organs  and  in  the 
heart,  which  act  as  a  suction-force  upon  it  and  constitute  the 
Tis  a  f route;  to  which  must  be  added  the  action  in  the  veins 
themselves,  which  are  muscular  and  richly  supplied  with 
nerves  connecting  with  the  vaso-motor  and  respiratory  centres. 
Well,  the  blood  in  the  arterial  system  of  warm-blooded  animals 
is  not  only  under  high  pressure, but  is  also  more  highly  oxygen- 
ated, being  of  a  bright  red  color,  the  two  going  together — the 
high  pressure  with  the  increase  in  oxygen.  So  that  we  have  an 
arrangement  for  increasing  circulation  in  the  tissues  with  an 
arrangement  for  increasing  metabolism,  the  two  being  in  cor- 
respondence. And  we  can  see  why  this  also  should  be  as  it 
is.  Since  it  relates  to  the  generation  of  force,  there  would 
be  nothing  gained  by  increasing  circulation  in  the  tissues  in 
the  absence  of  the  oxygen  ;  hence,  this  special  combination  for 
evolving  force  in  the  organism.  But  why  in  the  warm-blooded 
animal  m  ore  than  the  cold  %  This  also  is  easily  answered.  Thus, 
in  the  case  of  the  cold-blooded  animal  the  body  lies  prone  upon 
the  ground,  the  movements  are  sluggishly  and  awkwardly  per- 
formed, while  respiration  is  imperfect  and  digestion  slow  and 
delayed.  On  the  other  hand,  in  the  case  of  the  warm-blooded 
animal  the  body  is  suspended  off  the  ground  by  means  of  the 
crura,  which  involves  a  tremendous  struggle  with  gravitation, 
while  the  animal  is  in  constant  motion  and  the  movements 
energetic,  and  which,  of  course,  would  call  for  a  corresponding 


ARTERIALIZATION   OF   THE   BLOOD   AND   DEVELOPMENT.      41 

amount  of  force  for  effecting  these  actions  ;  hence  the  increase 
in  the  respiratory  and  digestive  functions,  together  with  the 
high  pressure  in  the  arterial  system  for  increasing  metabolism, 
for  in  no  other  way  could  it  be  done.  Again  everything  is  in 
correspondence.  * 

A  highly  interesting  and  important  fact  in  the  animal  cir- 
culation is  the  presence  of  iron  in  the  red  corpuscles  in  the 
form  of  haemoglobin,  for  increasing  polarity,  whereby  oxy- 
gen is  more  readily  pulled  into  the  organism  and  carbonic 
acid  expelled  from  it ;  since  it  is  not  a  chemical  union  which 
is  effected  with  these  gases,  but  one  in  which  the  molecules 
are  in  light  contact  simply,  the  haemoglobin  readily  yielding 
up  the  oxygen  in  the  tissues,  while  the  carbonic  acid  is  easily 
displaced  by  the  oxygen  when  the  venous  blood  is  exposed  to 
the  air.  And  by  reducing  pressure  simply,  as  in  a  receiver  and 
air-pump,  oxygen  escapes  from  the  corpuscles.  The  great  com- 
plexity in  this  albuminous  compound  (C  54.0,  H  7.25,  N  16.25, 
Fe  0  .42,  SO  .63,  O  21.45 — Hermann),  and  which,  of  course,  is 
necessary  for  producing  the  polar  conditions  spoken  of,  makes 
this  one  of  the  most  wonderful  adjustments  in  the  body.  But 
this  matter  is  more  advantageously  treated  in  connection  with 
development,  to  which  the  reader  is  referred,  the  more  im- 
mediate object  in  hand  being  to  show  the  law  in  the  circulation 
and  the  relations  it  sustains  to  pressure. 

Showing  that  the  Animal  Fluids  Respond  to  Changes  in 
Pressure. — Before  proceeding  to  make  application  of  the  pre- 
ceding principles  for  elucidating  vital  phenomena,  anatomical 
and  physiological,  it  might  not  be  amiss  to  call  to  mind 
familiar  circumstances  of  every-day  occurrence,  which  establish 
incontrovertibly  the  important  fact  that  the  animal  fluids  re- 
spond to  changes  in  pressure,  and  showing  conclusively  that  it 
is  fundamental  in  the  animal  organism,  thus  fully  preparing  the 
mind  for  what  is  to  follow  in  the  text.  For  this  purpose  we 
have  prepared  the  following  illustrative  diagram  (Fig.  8). 
For  example,  it  would  scarcely  be  contended  for  a  single  mo- 
ment even  that  the  principle  which  applies  for  aspirating  the 
secretions  in  the  mammary  gland  by  means  of  the  breast-pump 

*  For  the  rest  the  reader  is  referred  to  the  work  "  On  Gravitation  and  Develop- 
ment," where  many  other  matters  come  up  in  this  connection. 


42  PRESSURE   FUNDAMENTAL   IN   THE   BODY. 

(B)  is  not  identical  with  that  in  the  ordinary  syringe  (A),  the 
fluids  in  both  cases  flowing  into  the  instruments  by  reason  of 
the  low  pressure  which  is  developed  within  them  and  with 
which  they  communicate,  till  pressure  is  uniform.  Indeed, 
the  very  tissues  themselves  are  compressed,  pushed  and 
squeezed  into  the  breast-pump  in  order  to  equalize  press- 
ure, external  force  compelling  this  circumstance.  But  a 
yet  more  forcible  illustration  is  furnished  in  the  case  of 
the  ordinary  cuprjing-glass  (E).  In  this  contrivance  the  air  in 
the  cup  is  rarified  simply  by  the  burning  alcohol,  when  the 
instrument  is  suddenly  inverted  over  the  parts,  which  are  at 
once  forcibly  compressed  into  it,  the  blood  at  the  same  time 
flowing  rapidly  into  the  imprisoned  structures  in  order  to 
equalize  pressure,  till  the  distended  and  swollen  capillaries 
burst  with  their  contents,  producing  the  characteristic  ecchy- 
moses,  so  great  is  the  energy  in  this  force  when  suddenly  devel- 
oped. And  since  there  is  no  power  in  the  body  to  prevent 
this  abnormal  current  in  the  blood  and  juices,  it  follows  that 
pressure  is  the  fundamental  circumstance  in  the  organism 
with  which  everything  must  have  adjustment.  Indeed,  it 
furnishes  a  crucial  test  of  this  fact ;  while  the  actions  taking 
place  in  respiration,  in  the  heart,  arteries  and  hollow  viscera, 
are  to  be  interpreted  from  that  stand-point ;  otherwise  are 
meaningless. 

In  fine,  but  for  this  adjustment,  with  pressure  and  the 
power  of  producing  rapid  rhythmical  changes  in  pressure, 
animal  life  would  be  utterly  impossible. 

And  it  applies  not  only  to  respiration  and  the  actions  in  the 
heart  and  vessels  but  to  deglutition,  defecation,  urination — in 
short  to  all  the  hollow  viscera,  since  they  all  relate  to  circula- 
tion, and  the  maintenance  of  a  balance  in  the  organism ;  other- 
wise these  actions  also  would  be  meaningless.  Thus  in  the 
case  of  the  nursing  infant  (B)  can  it  be  doubted  for  a  single 
moment  even  that  the  same  principle  applies  for  compelling 
the  mammary  secretions  in  the  mouth-cavity,  as  in  the 
chamber  of  the  breast-pump  (B),  or  the  production  of  a 
lower  pressure  in  the  cavity  than  in  the  mammary  gland, 
the  fluid  flowing  from  high  to  low  pressure  in  conformity 
with  universal  law  \    (And  it  is  more  than  probable,  as   the 


PRESSURE  FUNDAMENTAL   IN   THE  BODY. 


43 


secretions  are  being  thus  withdrawn,  that  the  irritations  pro- 
pagated from  the  skin-surface  are  reflected  thence  upon  the 
milk  ducts,  which  causes  them  to  contract  for  increasing  press- 
ure, whereby  the  flow  into  the  mouth-cavity  is  expedited). 
Furthermore,  that  the  same  principle  applies  for  compelling 
them  thence  into  the  stomachal  cavity  represented  in  the 


Fig.  8.— Familiar  Modes  of  Pumping  the  Fluids  out  of  the  Body,  showing  that  they 
respond  to  changes  in  pressure. 

warns  of  deglutition,  which  consist  in  an  anterior  expansion 
for  diminishing  pressure  with  a  posterior  contraction  for  in- 
creasing it,  whereby  the  journey  of  the  fluid  or  the  bolus  along 
the  canal  is  greatly  expedited,  the  waves  following  each  other 
in  rapid  succession  till  the  stomachal  cavity  is  reached.     In 


44  PRESSUBE   FUNDAMENTAL   IN   THE   BODY. 

short,  that  deglutition  is  based  upon  rhythmical  changes  in 
pressure,  otherwise  is  inexplicable.  Also,  that  the  same  applies 
for  the  sucking  leech  (c),  which  is  feeding  in  the  capillaries  of 
its  victim,  while  the  undulations  coursing  along  its  tubular 
body  from  the  mouth-cavity  are  analogous  with  deglutition,  of 
which  it  is  archetypal,  the  principle  in  mechanics  being  the 
same  for  both,  while  the  structures  are  fundamentally  the 
same,  of  which  more  anon. 

Finally,  that  pressure  is  transmitted  through  the  body  upon 
all  the  organs  is  also  of  easy  demonstration.  Notably  the  in- 
strument of  Dulafoy  for  aspirating  depots  of  pus  and  collections 
in  the  deep  tissues  and  organs  of  the  body,  furnishes  a  crucial 
test  of  this  circumstance.  For  example,  it  establishes  the  im- 
portant fact  that  an  empty  air-tight  vessel  when  placed  at  the 
bedside  of  a  patient,  and  made  to  communicate  with  an  internal 
collection  by  means  of  an  elastic  tube  and  canula,  will  aspirate 
the  collections  the  moment  that  pressure  is  reduced  within  the 
instrument  by  pumping  out  some  of  the  air  it  contains,  till  the 
very  last  portions  are  removed.  Furthermore,  that  it  will 
do  this  at  a  distance  from  the  body;  this  in  the  absence  of  any 
other  means  for  assisting  it  through  the  tubing,  the  difference 
in  pressure  in  the  two  localities  being  sufficient  for  the  purpose. 

The  practical  deduction  to  be  made  from  this  circumstance 
is,  that  the  extensive  arrangements  which  obtain  for  reducing 
pressure  in  the  lungs  during  inspiration  should  have  the  effect 
of  aspirating  the  venous  system  at  the  same  time  that  it  aspi- 
rates air,  while  the  action  in  the  heart  should  greatly  expedite 
it,  whereby  correspondence  is  produced  between  the  circula- 
tion of  air  and  blood  in  the  alveoli,  as  before  remarked.  It  is 
needless  to  extend  the  matter. 

In  conclusion,  the  power  of  producing  the  rhythmical  expan- 
sions and  contractions  for  effecting  the  requisite  changes  in 
pressure  in  the  organs  and  tissues  for  expediting  circulation 
and  producing  the  voluntary  movements  (the  principle 
is  the  same  in  both)  inheres  in  protoplasm  itself,  which 
expands  and  contracts  with  great  facility ;  moreover,  expan- 
sion and  contraction  are  correlated  forces  in  Nature.  The 
molecular  action  it  involves  will  come  up  at  the  proper  time 
and  place.     (See  closing  chapter.)    Hoping  we  have  made  our- 


CONCLUSION   OF  PEINCIPLES.  45 

selves  understood  sufficiently,  let  us  now  turn  our  attention  to 
the  circumstances  in  development  explanatory  of  this  funda- 
mental principle  in  the  mechanics  of  circulation,  taking  the 
vital  phenemena,  anatomical  and  physiological,  as  the  text, 
and  the  law  of  pressure  underlying  the  organism  as  the  key 
for  construing  and  interpreting  them,  or  as  means  to  ends 
simply.  And  since  there  is  increasing  differentiation  in  the 
organs  and  tissues  with  progress  in  development,  for  the  sake 
of  simplicity  and  effectiveness,  it  were  best  to  begin  with  the 
lowest  organisms,  taking  the  action  in  undifferentiated  proto- 
plasm itself  as  the  first  visible  expression,  proceeding  thence 
as  rapidly  as  possible  till  the  highest  forms  are  reached. 
Indeed,  one  must  proceed  from  the  simpler  to  the  more  com- 
plex forms  in  order  to  make  the  latter  intelligible.  In  this 
manner  the  whole  is  illuminated,  and  everything  made  plain 
and  easily  understood. 


CHAPTER  IV. 

RESPIRATION  IN   DIFFERENT   STAGES   IN  DEVELOPMENT. 

Import  of  Amoebae  Movement — The  Alternate  Extension  and  Retraction  of  the  Branched 
'  Processes,  a  Pumping  Action  for  Increasing  Circulation — Why  Locomotion  Should 
Increase  Circulation  Correspondingly — The  Pumping  Movements  Analogous  with 
Respiration — The  Action  in  Vacuoles  and  the  Radiating  Canals,  an  Early  Indication 
of  the  Mechanical  Principle  in  the  Heart  and  Arteries,  the  Latter  Expanding  as  the 
Former  is  Contracting,  and  vice  versa — The  Action  in  Gastrula — Necessity  for  Co- 
ordinating the  Mucous  with  the  Skin-Surface,  in  Order  to  Produce  Afflux  and 
Efflux  of  the  Fluids  in  the  Body-Interior— This  Circumstance  Further  Illustrated  in 
the  Worms  for  Producing  the  Undulations  which  Course  Along  the  Body  During 
Imbibition — The  Principle  Applied  to  Respiration  and  the  Action  Taking  Place  in 
the  Lungs — Illustrated  in  the  Frog,  in  which  it  is  Demonstrated  Experimentally 
that  the  Lungs  Expand  and  Contract  Regularly  and  Rhythmically  Synchronous 
with  the  Action  Taking  Place  in  the  Muscular  Envelope  or  Containing  Walls,  in 
Order  to  Produce  Afflux  and  Efflux  of  Air  and  Blood  in  these  Organs  for  Respira- 
tory Purposes  ;  Otherwise  Impossible — The  Manner  the  Parts  are  Coordinated — 
Dependence  of  the  Portal  Circulation  upon  Respiration— The  Same  Principle 
in  Mechanics  for  Every  Stage  in  Development— The  Action  in  Birds— The  Special 
Adjustments  in  the  Viscera— Portal  Circulation  in. 

This  relation  which  animal  life  sustains  to  pressure,  and 
the  law  for  increasing  circulation  by  rapid  rhythmical  changes 
in  £>ressure,  would  afford  a  ready  explanation  for  the  alternate 
extension  and  retraction  of  the  branched  processes  in  amoebae, 
and  which  function  as  temporary  villi  for  pumping  the  com- 
merce in  the  organism.  For  example,  when  an  amoeba  is 
placed  upon  the  slide  in  the  field  of  the  microscope,  and  a 
drop  of  water  suffered  to  fall  upon  it,  it  at  once  begins  to 
extend  and  retract  the  branched  processes  in  rapid  succession, 
and  in  every  direction  from  the  undifferentiated  body,  appear- 
ing and  disappearing,  now  here,  now  there,  in  the  protoplasmic 
substance.  It  is  not  trying  to  get  away,  but  trying  rather  to 
live  and  sustain  existence  by  means  of  these  branched  pro- 
cesses and  this  pumping  action  thus  set  up  in  them.  In  short, 
the  animal  is  simply  feeding ;  nay,  feeding  and  respiring  at 
the  same  time,   the  one  involving    the    other ;    and  which 


PUMPING  ACTION   IN   AM(EB^E.  47 

should  be  the  case  for  maintaining  a  balance  in  force,  the  same 
principle  for  generating  force  applying  for  it  as  for  the  higher 
stages  in  development,  and  must  necessarily  apply  for  every 
stage.  These  so-called  pseudopodia  (false  feet),  then,  are  not 
for  effecting  locomotion  simply,  as  their  name  implies,  but  for 
sustaining  existence  ;  at  the  same  time,  they  are  available  for 
locomotion,  though  this  is  laboriously  and  indifferently  per- 
formed. Furthermore,  it  is  readily  perceived  that  this  also 
should  increase  circulation  correspondingly,  since  it  involves 
more  extended  and  energetic  action  in  the  branched  processes 
for  effecting  it,  whereby  the  requisite  force  is  generated  for 
producing  this  action  ;  otherwise  impossible.  In  this  manner 
a  balance  is  maintained,  one  adjustment  involving  the  other. 

In  fine,  when  the  amoeba  extends  a  branched  process,  it 
develops  a  lower  pressure  within  itself  than  exists  externally, 
when  the  fluids  immediately  adjacent  rush  into  the  low- 
pressure  area  until  pressure  is  uniform,  and  which  corresponds 
with  inspiration.  But  when  the  branched  processes  are 
retracted,  this  compels  the  contents  into  the  body -interior, 
at  the  same  time  expelling  waste  products,  and  which  corre- 
sponds with  expiration.  Of  course,  the  more  rapid  the  action 
the  more  rapid  is  this  circulation.  The  in-going  current  con- 
tains the  nutritive  and  force -producing  elements,  and  the  out- 
going the  waste  products  ;  while  water  is  the  vehicle  or 
medium  of  transportation  for  the  commerce,  at  the  same  time 
it  enters  into  the  molecular  arrangements  for  forming  the 
structure  itself.  In  addition,  water  is  a  powerful  stimulus  for 
exciting  the  pumping  actions  and  arousing  latent  energy,  so 
that  the  moment  it  touches  the  amoebae  it  begins  to  pump,  and 
thus  compelling  it  into  itself,  the  whole  mechanism  becomes 
"alive." 

But  remove  the  water,  and  the  pumping  action  stops  and 
life  is  in  abeyance.  No  one  can  tell  what  water  is — extend- 
ing as  it  does  into  the  depths  of  force — further  than  to  speak 
of  some  of  its  performances  and  the  conditions  which  deter- 
mine them,  notably  the  polar  actions  and  the  means  of  ener- 
gizing them,  inclusive  of  the  law  of  pressure  for  increasing  the 
action  in  the  manner  spoken  of,  or  by  rhythmical  changes  in 
pressure  for  compelling  it  in  and  through  the  organism  in  the 


48  PUMPING  ACTION   IN   AMCEB^E. 

measure  of  the  requirements.  Furthermore,  by  extending 
and  retracting  the  branched  processes  from  different  portions 
of  the  body-surface,  the  amoeba  is  enabled  to  effect  a  short 
and  direct  journey  of  the  fluids  to  the  portions  where  they  are 
most  needed,  and  which  subserves  the  purpose  of  a  vascular 
system  for  effecting  the  same  ends  in  the  higher  stages  in 
development ;  at  the  same  time,  it  also  involves  an  amount  of 
motion  of  the  fluids  loitliin  the  body,  whereby  metabolism  is 
expedited.  Thus,  when  a  branch  process  is  being  extended, 
the  low  pressure  which  is  thereby  developed  within  this  por- 
tion produces  afflux  of  the  fluids  in  two  directions — one  from 
the  body  itself,  the  other  from  the  environment — coming  from 
opposite  directions  simultaneously  to  the  low-pressure  area 
till  pressure  is  uniform ;  while  in  the  body  itself  a  special 
force  applies  for  increasing  the  action,  notably  the  contraction 
which  sets  in  simultaneously  with  extension  in  the  branched 
process,  should  effect  a  corresponding  increase  in  pressure 
for  compelling  the  contents  into  the  branched  process,  the 
body  fluids  flowing  into  this  more  rapidly  in  consequence. 
On  the  other  hand,  when  the  process  is  being  retracted  there 
is  commensurate  expansion  in  the  body  in  order  to  effect  it, 
the  former  producing  high,  the  latter  low  pressure  ;  hence,  the 
fluids  must  flow,  out  of  the  branched  process  into  the  body- 
interior  with  corresponding  energy.  Thus,  a  churning  move- 
ment is  produced  in  the  fluids  for  hastening  metabolism.  The 
following  forcible  illustration  by  the  distinguished  biologist 
at  Heidelberg  (Fig.  7,  A,  B)  will  serve  for  impressing  the  mat- 
ter. It  represents  an  amoeba  at  two  different  moments  during 
movement.  By  fixing  the  eye  upon  the  food  particle,  we  can 
very  readily  perceive  how  the  contents  are  affected  by  the 
pumping  action  in  the  processes.  Thus,  in  the  case  of  B,  in 
which  a  branched  process  is  extended,  the  body  is  narrow  or 
contracted,  while  the  food  particle  is  moved  down  to  the  root 
of  the  process ;  but  in  A,  in  which  the  process  is  retracted, 
the  body  is  considerably  widened  or  expanded,  and  now  the 
food  particle  (/)  is  moved  up  to  near  the  central  portions  of 
the  body.  Thus,  a  to-and-fro  movement  is  established  be- 
tween the  fluids  in  the  body  and  the  branched  processes, 
thereby  effecting  more  rapid  disintegration  and  assimilation 


THE   INCREASE   OF   RESPIRATION  AND   CIRCULATION.         49 

of  the  food,  at  the  same  time  that  it  pumps  the  fluids  in  and  out 
of  the  body  for  respiratory  purposes,  inclusive  of  the  nutritive 
elements  in  a  state  of  solution  Hence  the  rapid  absorption 
of  tine  coloring  matter,  which  passes  into  the  amoeba  with 
great  rapidity. 

Finally,  that  this  pumping  action  in  amoeba  relates  to  the 
nutritive  and  functional  processes  is  sufficiently  obvious,  there 
being  no  other  means  for  increasing  circulation.  Moreover, 
the  same  action  is  seen  in  Bathybius — enormous  masses  of 
retiform  protoplasmic  substance  covering  the  floor  of  the 
ocean,  in  places  to  many  feet  in  depth — which  does  not  loco- 
mote  nevertheless  performs  similar  movements  when  a  frag- 
ment is  placed  in  the  field  of  the  microscope,  and  a  drop  of 
water  is  suffered  to  fall  upon  it,  the  action  undoubtedly  indicat- 
ing respiratory   movement  for  increasing  circulation  in  the 


Fig.  9. — An  Amoeba  figured  at  two  different  moments  during  movement ;  n,  nucleus , 
i,  ingested  food.     Some  vacuoles  may  also  be  noted. — Gegenbaur. 

protoplasmic  substance  and  interstices,  the  surrounding  fluids 
flowing  in  and  out  through  these. 

Modes  of  Introducing  Solid  Food  in  Amosbce. — The  me- 
chanical principle  which  applies  for  absorbing  solid  food  in 
amoebae  is  the  same  as  for  producing  the  pumping  actions, 
notably  by  expanding  and  contracting  upon  it,  only  it  has 
different  expression,  and  which  varies  with  the  stage  in 
development.  Thus,  in  the  lowest  amoebae,  in  which  the  body- 
surface  is  naked  and  undifferentiated  protoplasm,  solid  food  is 
absorbed  in  two  different  ways  :  either  by  extending  branched 
processes  around  the  solid  particle,  when  the  soft  substance 
flows  together  again,  and  so  engulfing  it ;  or  else  by  the  body 
itself  opening  at  the  point  of  contact  for  receiving  it,  the  food  at 


50  INTRODUCTION   OF   SOLID   FOOD. 

the  same  time  sinking  into  it  from  the  action  of  external  press- 
ure till  fully  submerged,  when  the  substance  closes  over  it. 
In  either  case  involving  expansile  and  contractile  action  in  the 
protoplasm  in  order  to  effect  it ;  while  this  in  turn  is  referable 
to  the  action  in  the  molecules,  and  which,  of  course,  is  regu- 
lated as  occasion  may  require.  And  any  portion  of  the 
body-surface  may  thus  function  as  a  mouth  or  receiving  organ 
for  the  food,  just  as  any  portion  may  serve  for  extending  and 
retracting  the  branched  processes  for  pumping  in  the  fluids, 
while  pressure  applies  alike  to  both  actions  for  making  them 
effective.  Thus,  when  the  surface  opens  in  response  to  the 
stimulus  of  the  food  brought  into  contact  with  it,  a  suction 
force  is  at  once  produced  for  compelling  it  into  the  organism, 
and  which  necessarily  results  from  the  act  of  expansion,  the 
particle  sinking  deeper  and  deeper  with  progress  in  expansion, 
till  finally  it  reaches  the  locality  where  it  is  to  undergo  diges- 
tion and  assimilation,  there  being  no  stomach  or  digestive 
cavity  for  receiving  it  when  the  protoplasmic  substance  closes 
over  it.  Likewise,  when  a  cortical  layer  and  external  organs 
are  differentiated  (Fig.  10),  the  same  principle  applies. 

In  this  case  the  radiating  processes  (e)  serve  as  tactile  and 
prehensile  organs  for  directing  the  food  to  the  part  which  is 
to  function  at  the  time  as  a  mouth  or  receiving  organ,  holding 
it  against  the  surface  (a)  till  the  requisite  changes  can  be  made 
in  the  adjacent  parts,  when  it  is  compelled  into  the  body  under 
the  action  of  external  pressure,  where  their  function  ends  in 
this  respect.  And  let  it  not  be  thought  for  a  single  moment 
even  that  they  can  compel  the  food  into  the  body,  for  this  rude 
mechanics  is  not  only  wrong  in  principle,  but  must  inevitably 
prove  disastrous,  since  the  action  necessarily  involves  appro- 
priate internal  arrangements  for  receiving  the  food,  while  the 
parts  yield  of  their  own  accord  under  the  stimulus  of  the 
food,  a  change  at  once  setting  in  at  the  point  of  contact,  and 
extending  thence  to  the  adjacent  protoplasm,  which  in  like 
manner  recedes  before  it  till  it  reaches  its  destination,  all 
the  parts  acting  in  harmonious  concert  to  this  end.  In 
other  words,  room  is  made  for  the  food  as  it  progresses 
into  the  body,  while  the  very  effort  to  make  this  room  in- 
vokes the  force  of  external  pressure  for  compelling  it  into 


INTRODUCTION   OF   SOLID   FOOD. 


51 


the  body.  And  from  the  moment  it  starts  upon  the  journey 
till  it  reaches  its  destination,  such  is  the  case.  And  one  morsel 
after  another  being  thus  taken  in,  when  the  amoeba  is  made 
to  eat,  the  limit  in  expansion  is  finally  reached  when  no  more 
food  can  be  taken  into  the  body,  a  circumstance,  indeed,  which 
is  forcibly  illustrated  in  a  higher  stage  in  development,  e.  g. , 
the  gorged  leech  (Pig.  17).  But  we  will  not  anticipate. 
When  there  is  a  mouth,  however  (Ciliata),  this  is  expanded 
when  brought  in  contact  with  the  food,  pressure,  of  course, 
forcing  it  in  simultaneously,  whence  it  passes  into  the  soft 
parenchyma  as  previously,  as  there  is  no  enteric  tube  for  con- 
veying the  food  masses,  the  parts  again  contracting  and  closing 
over  it.  Indeed,  the  powers  of  expansion  possessed  by  these 
low  organisms  is  something  extraordinary,  some  of  the  more 
ravenous  infusoria,  for  example,  actually  swallowing  other 


Fig.  10.— Actinosphserum  (Gegenbaur).  a,  A  morsel  which  has  been  taken  in  as  food, 
and  just  pushed  into  the  soft  cortical  layer  b,  by  a  change  in  pressure ;  c,  central 
parenchyma  ;  d,  other  food  which  had  previously  been  thus  introduced  ;  e,  pseudo- 
podia  of  the  cortical  layer. 

infusoria  nearly  as  large  as  themselves.  The  mouth  being 
brought  in  contact  with  the  prey,  is  suddenly  expanded  upon 
it,  which  at  once  develops  a  suction  force  for  compelling  it  into 
the  cavity,  when  the  mouth  contracts  again  for  completing 
the  act.  Other  infusoria  (Suctoria)  possess  hollow  radiate 
processes  (Fig.  11,  p.  55),  which  pass  through  the  envelope  of  the 
body  and  function  as  suckers,  so  to  speak,  and  which  are 
made  to  penetrate  the  body  of  other  infusoria  for  aspirating 
the  fluids  into  themselves,  flowing  into  them  in  the  form  of 
drops,  thus  feeding  upon  the  juices  of  the  victim  simply,  not 
possessing  a  mouth.    But  for  aspirating  the  fluids,  of  course 


52  CARDIAC   ACTION   IN   VACUOLES. 

the  body  must  expand  for  reducing  pressure,  the  fluids  in 
consequence  flowing  into  themselves  from  the  body  of  the 
victim  where  pressure  is  higher. 

Finally,  since  there  is  no  means  for  compelling  food  into  the 
body  save  by  changes  in  pressure,  it  follows  that  but  for  this 
power  of  expanding  and  contracting,  which  inheres  in  proto- 
plasm, that  the  food  must  inevitably  remain  out  of  the  body,  for 
in  no  other  way  could  the  requisite  changes  in  pressure  be 
produced.     In  fine,  that  animal  life  would  be  impossible. 

The  Action  in  Vacuoles  and  Wliat  it  Involves. — The  hol- 
low cavitary  spaces  filled  with  a  colorless  fluid  and  known  as 
vacuoles  (Fig.  11,  v,  v),  is  the  first  foreshadowing  of  a  spe- 
cialized vascular  apparatus  for  distributing  the  fluids  through 
the  body,  while  the  phenomena  manifested-  in  them  possess 
special  significance  with  reference  to  the  principle  that  obtains 
in  the  heart  and  arteries.  Thus,  the  vacuole  expands  and 
contracts  regularly  and  rhythmically  like  the  diastole  and 
systole  in  the  heart,  while  a  system  of  radiating  canals  (at 
present  invisible)  receive  and  discharge  the  fluids  from  the 
pulsating  vacuole,  expanding  and  contracting  alternately  with 
it,  so  that  during  systole  in  the  vacuole  there  is  diastole  in 
the  radiating  canals,  and  vice  versa  ;  the  vacuole  itself,  like 
the  radiating  canals,  coming  into  view  during  diastole,  from 
the  presence  of  the  fluids,  which  give  it  definition,  and  dis- 
appearing again  with  systole  when  the  fluids  are  in  the  radi- 
ating canals,  the  two  alternately  appearing  and  disappearing 
in  this  manner  and  in  regular  order  and  succession,  while  the 
fluids  flow  from  high  to  low  pressure  in  conformity  with  uni- 
versal law.  The  following  lengthy  excerpt,  however,  will 
place  the  matter  fully  before  the  reader  : 

*  k '  More  definite  respiratory  arrangements  are  seen  when,  as  in 
many  protozoa,  water  is  taken  into  the  body.  Cavities,  which 
are  filled  with  a  fluid,  and  which  gradually  contract  and  com- 
pletely empty  themselves,  after  having  reached  their  maxi- 
mum of  distension,  appear  within  the  protoplasm ;  when  empty 
they  seem  to  disappear.  These  vacuoles,  like  the  vacuoles  in 
the  cells  of  certain  tissues,  are  partly  variable  structures,  now 

*  Elements  of  Comparative  Anatomy,  pp.  85.  86— Gegenbaur. 


PRIMITIVE   CIRCULATION.  53 

appearing  and  now  disappearing,  and  partly  constant.  When 
they  are  constant,  their  function  is  increased,  and  they  often 
expand  and  contract  regularly  and  rhythmically,  like  the  car- 
diac systole  and  diastole.  Contractile  vesicles  of  this  kind  are 
often  seen  in  the  amoebae  (Difflugia  and  Arcella),  and  are  very 
common  among  the  infusoria.  They  are  also  known  as  vacu- 
oles. The  fluid  which  collects  in  the  vesicles  is  drawn  from 
the  parenchyma  of  the  body  and  is  ieturned  to  it,  or  passed 
out  to  the  exterior  on  the  contraction  of  the  vesicle  Fine 
communications  with  the  exterior  have  been  made  out,  so  that 
the  latter  course  is  the  probable  one  ;  but  we  need  not,  on  this 
account,  conclude  that  water  does  not  enter  by  the  same 
passage. 

' k  In  the  infusoria  the  vesicles  lie  in  the  cortical  layer,  generally 
just  under  the  delicate  cuticle,  and  at  definite  points.  If  only 
one  vesicle  is  present,  it  lies  either  anteriorly  or  posteriorly ; 
if  two,  there  is  one  near  each  end  of  the  body.  Trachelius 
ovum  is  remarkable  for  a  large  number  of  small  vesicles.  No 
special  membranes  can  be  made  out  on  the  wall  of  the  vesicle 
nor  in  the  canals  which  pass  off  from  it.  Like  the  vesicle,  the 
canals  can  only  be  made  out  while  they  are  filling.  The  vesicle 
and  canals  contract  alternately.  In  Paramecium  the  canals 
enlarge  at  the  commencement  of  the  systole  and  approach  one 
another  as  the  vesicle  diminishes  in  size,  so  that  they  form  a 
stellate  figure  at  the  moment  when  its  systole  is  most  complete 
and  the  vesicle  has  disappeared.  While  the  vesicle  is  filling, 
the  canals  look  like  small  diverticula  on  it,  and  are  not  again 
fully  distended  until  the  diastole  is  complete.  The  number  of 
the  canals,  which  is  limited  in  P.  aurelia  to  eight  or  ten,  is 
increased  to  thirty  in  Bursaria  flava,  and  is  much  higher  in 
Cystostomum  leucas.  In  these  forms  the  canals  have  a  wave- 
like course,  and  ramify  at  their  extremities  Canalicular 
tracts  are  formed  by  the  fusion  of  several  spaces  filled  with 
water  into  longer  tracts,  as  in  Stylonychia  (St.  mytilus),  and 
they  empty  themselves  into  the  contractile  vesicle  by  definite 
passages.  The  long  canals  of  Spirostomum  ambiguum,  which 
are  also  visible  for  a  time  only,  but  which  are  longer  than 
these,  are  like  them,  so  that  we  can  make  out  a  continuous 
series  from  the  first  appearance  of  an  apparently  indifferent 


51     ACTION  IN  THE  HEART  AND  ARTERIES  FORESHADOWED. 

cavity  to  a  definitely  arranged  system  of  tubes."  Italics 
are  added.  There  can  be  no  doubt,  then,  that  the  func- 
tion of  vacuoles  relates  to  the  distribution  of  the  fluids 
through  the  body,  and  which  involves  not  only  respiration 
and  expulsion  of  waste  products,  but  the  nutritive  proc- 
esses as  well,  the  fluids  containing  both  nutritive  and  force- 
producing  elements.  But  the  point  we  wish  to  make,  how- 
ever, concerns  the  mechanical  principle  which  applies  for 
filling  and  emptying  the  vacuole  and  radiating  canals,  and 
which  is  undoubtedly  rhythmical  changes  in  pressure.  Thus 
when  the  vacuole  expands  it  develops  a  lower  pressure 
within  itself  than  exists  externally :  hence  the  fluids  from 
the  adjacent  parenchyma  and  environment  (Fig.  11,  v,  v.) 
flow  into  the  low-pressure  area  till  pressure  is  uniform  ;  but 
when  contraction  sets  in,  the  high  pressure  which  this  develops 
within  the  vacuole  causes  them  to  flow  out  again,  passing 
thence  into  the  radiating  canals,  which  expand  simultaneously 
for  reducing  pressure  and  so  obviating  reflux  ;  thence  into  the 
environment  during  contraction  and  development  of  high 
pressure  in  the  latter.  And  so  by  alternate  expansions  and 
contractions  in  the  vacuole  and  radiating  canals,  the  fluids  are 
pumped  through  the  body  for  respiratory  purposes ;  other- 
wise is  meaningless.  In  short,  it  represents  a  suction  and  a 
driving  force,  acting  alternately ;  the  one  produced  by 
diastole,  the  other  by  systole.  The  eminent  anatomist  and 
naturalist  would  seem  to  emphasize  systole,  but  the  fluids 
would  first  have  to  be  gotten  into  the  vesicle  by  diastole, 
and  the  development  of  low  pressure  which  this  produces, 
before  one  may  speak  of  systole  for  driving  them  out  again  ; 
the  one  necessarily  involving  the  other.  But  exclude  press- 
ure and  the  power  of  producing  rhythmical  changes  in  press- 
ure, and  these  actions  would  be  meaningless.  Now,  then, 
principles  do  not  and  cannot  change  ;  hence  this  law  for 
increasing  circulation  remains  the  same  for  every  stage  in 
development.  And  in  this  primitive  circulation,  in  which  not 
so  much  as  a  living  membrane  is  made  out,  either  in  the  vacu- 
ole itself  or  the  radiating  canals,  which  are  wrought,  as  it 
were,  in  naked  and  undifferentiated  protoplasm,  the  principle 
in  the  heart  and  arteries  is  clearly  indicated  ;  the  one  expand- 


ACTION  IN  THE  HEAET  AND  AKTEEIES  FORESHADOWED.      55 


ing  as  the  other  is  contracting,  and  vice  versa;  while  the 
valves  serve  for  obviating  reflux  ;  and  leaving  not  the  shadow 
of  a  doubt  upon  the  mind  of  the  power  in  Nature  to  effect 
these  adjustments. 

But  when  we  come  to  the  blood-vascular  system,  how- 
ever, ample  proof  will  be  furnished  of  this  circumstance. 
Suffice  it  to  say,  the  law  applies  everywhere.  And  the  more 
one  reflects  over  this  circumstance,  the  more  wonderful  it 
grows.  This  will  serve  for  illustration.  And  we  will  now 
carry  the  matter  a  little  further,  pushing  the  law  in  the  organ- 
ism to  its  logical  results. 


Eig.  11. — An  Acineta,  with  part  of  its  stalk  (Gegenbaur)  ;  p,  pseudopodia-like,  but  stiff, 
tentacles  ;  v,  vacuoles  ;  n,  nucleus  ;  e,  aciliated  young  form  lying  in  the  so-called 
broad  cavity. — G-egenbaur. 

The  Action  in  Gastrula. — This  brings  us  to  the  next  stage 
in  development,  or  the  primitive  compound  organism  (Fig.  12), 
from  which  in  due  time  the  higher  organisms  are  evolved  by 
the  processes  of  growth  and  differentiation.  Here  the  cell- 
colony  is  first  grouped  in  two  separate  and  distinct  layers, 
forming  the  walls  of  the  little  animal  (B),  the  one  constituting 
the  internal  layer,  or  entoderm  (i),  the  other  the  skin-layer,  or 
exoderm  (e);  while  the  fluids  flow  in  and  out  of  the  little 
stomachal  cavity  thus  formed  through  the  oral  orifice  (o)  dur- 
ing the  rhythmical  expansions  and  contractions  taking  place 
in  it.     Flowing  in,  of  course,  during  expansion  and  the  devel- 


56  ACTION    IN   GASTRULA. 

opment  of  low  pressure  which  this  produces  till  pressure  is 
uniform,  and  flowing  out  again  by  reflux  action  through  the 
route  of  ingress  during  contraction  and  the  development  of 
high  pressure  which  this  produces  to  equalize  pressure  ;  in  this 
manner  readily  producing  afflux  and  efflux  of  the  fluids  in  the 
stomachal  cavity  :  while  the  changes  of  form  which  this  in- 
volves in  the  individual  cells,  in  order  to  produce  these  actions 
should  increase  circulation  in  them  correspondingly,  and  which 
the  scheme  calls  for  in  order  to  generate  the  force  which  is 
expended  in  these  actions,  the  one  involving  the  other.     .All of 


Fig.  12—  Gastrula  of  a  Chalk-sponge  (Olynthus).  A,  from  the  outside  ;  B,  in  longi- 
tudinal section  through  the  axis ;  g,  primitive  intestine  ;  o,  primitive  mouth  ;  i, 
intestinal-layer,  or  entoderm  ;  e,  skin-layer,  or  exoderm. — Haeckel. 

which  is  plain  enough.  But  the  circumstance  in  which 
interest  culminates  is  the  one  of  coordination  in  the  cell-colony, 
so  as  to  compel  the  two  layers  to  expand  and  contract 
simultaneously,  in  order  to  swallow  the  food  and  effect  the 
pumping  action  for  producing  afflux  and  efflux  of  the  fluids  in 
the  primitive  intestine  (g) ;  otherwise  impossible.  In  short,  the 
inner  layer  (/)  or  intestine  expands  and  contracts  snychronously 
with  the  outer  wall  or  skin-layer  (e),  which  answers  to  the 
muscular  envelope  in  the  body- walls  in  the  more  advanced 
stages  in  development,  the  two  acting  in  harmonious  concert 
to  this  end,  as  is  the  case  in  the  later  stages  ;  only  there  is  no 
nervous  apparatus  for  effecting  coordination,  the  walls  expand- 


COORDINATION   OF   INTERNAL   AND   EXTERNAL   PARTS.        57 

Ing  and  contracting  of  their  own  accord  under  the  stimnlus  of 
the  fluids  in  which  the  little  animal  swims  and  sustains  exist- 
ence. The  intimate  relations  the  walls  sustain  to  each  other 
enable  this  to  be  done,  as  the  inter-molecular  action  in  the 
two  cell-layers  could  thus  be  connected  for  producing  uni- 
formity throughout.  But  with  progress  in  development,  how- 
ever, the  layers  become  more  and  more  differentiated  and 
separated  from  each  other ;  consequently,  other  modes  for  effect- 
ing coordination  must  be  had  recourse  to  ;  hence  the  nervous 
apparatus  which  is  developed  and  differentiated  with  progress 
in  development  for  coordinating  the  structures  and  unifying 
the  molecular  actions,  to  the  end  that  a  balance  in  force  may  be 
maintained,  and  which  cannot  be  done  unless  the  mechanics  is 
made  to  connect  through  and  through.  It  is  comprehensive, 
but  easily  understood.  J  n  this  archetypal  form,  then,  we  have 
early  indication  of  the  principle  in  the  mechanics. 

Indeed,  the  very  existence  of  this  law  of  pressure  in  the 
organism  is  sufficient  in  itself  to  establish  that  circumstance, 
now  that  attention  is  directed  to  it.  But  a  few  pointed  exam- 
ples will  serve  for  impressing  the  matter. 

Leaving  the  Gastrula,  then,  we  take  up  the  case  in  the 
"Worms,  in  which  development  is  considerably  advanced. 
Here  the  external  layer  (exoderm)  is  differentiated  into  a 
dermo-muscular  layer,  and  the  internal  (entoderm)  into  the 
intestine  (Fig.  1 3,  A,  B,  m,  v)  ;  but  are  still  more  or  less  inti- 
mately connected  by  means  of  connective  tissue  fibres  running 
from  one  to  the  other,  a  coelon  or  visceral  cavity  not  yet  be- 
ing differentiated,  the  vestiges  only  presenting  in  rudimentary 
spaces  adjacent  to  the  intestine,  in  which  the  fluids  collect  and 
pass  into  the  vascular  system.  For  producing  and  coordinat- 
ing the  movements  in  the  two  surfaces,  and  maintaining  uni- 
formity, we  have  a  double  ganglionic  chain  of  nerves  extend- 
ing along  the  ventral  surface  from  the  cephalic  to  the  caudal 
end  (Fig.  14),  each  ganglion  serving  as  a  separate  centre  of 
nervous  force  for  producing  the  movements  in  the  contiguous 
parts  into  which  the  nerves  are  distributed,  so  that  reflex 
action  is  readily  excited. in  them  by  means  of  sensory  impres- 
sions propagated  from  the  mucous  surface  by  the  stimulus  of 
the  food,  without  involving  other  portions,  and  which  is  neces- 


58       COORDINATION   OF  INTERNAL   AND   EXTERNAL   PARTS. 

sary  in  order  to  produce  the  undulations  passing  along  the 
body  of  the  worms  during  imbibition  and  the  passage  of  food 
along  the  canal  ;  at  the  same  time,  the  whole  are  readily  co- 
ordinated by  means  of  the  nervous  links  or  commissures 
extending  from  ganglion  to  ganglion  for  connecting  them  with 
each  other  and  with  the  encephalon  (Fig.  14)  for  producing  the 
voluntary  movements  and  maintaining  a  balance  in  the  organ- 
ism. There  is  considerable  complexity,  arising  from  the  neces- 
sity of  maintaining  the  local  actions  in  correspondence  with 
the  physiological  requirements,  but  the  principle  underlying 
it  all  is  readily  apprehended. 

Now,  then,  in  order  to  produce  the  undulations  which  course 
along  the  body  of  the  worms  during  imbibition,  and  which 
answer  to  the  passage  of  the  food  along  the  intestinal  canal, 


Fig.  13. — Transverse  section  of  Asoaris  Luinbricoides,  A,  and  of  Hirudo,  B.  c,  Cuticular 
layer  ;  to,  muscular  layer  ;  r,  lateral  line,  with  the  excretory  organ  ;  p  p,  upper  and 
lower  median  line  :  p1,  oblique  fibres  ;  v,  enteron  ;  d,  dorsal  ;  I,  lateral  vascular 
trunk  ;  s,  vesicle  of  the  excretory  organ  ;  n,  ventral  nerve-chord. — Gegenbaur. 

doth  of  these  layers  must  expand  and  contract  together  and 
simultaneously,  as  in  gastrula,  for  pumping  the  fluids  into  and 
out  of  the  cavity  of  the  intestine  ;  only  that  in  the  present  case 
this  has  been  prolonged  into  a  canal,  but  for  obvious  reasons 
the  principle  is  maintained,  since  the  whole  relates  to  pressure 
and  the  power  of  producing  rapid  rhythmical  changes  in 
pressure.  In  fine,  the  worms  may  be  likened  to  a  string  of 
gastrulse  with  the  ends  merged  in  each  other,  while  the  fluids 
are  passed  from  one  to  the  other  by  means  of  rhythmical 
expansions  and  contractions  till  the  terminal  end  is  reached, 


COORDINATION   OF  INTERNAL   AND   EXTERNAL   PARTS.        59 

every  pair  of  nervous  ganglia,  with,  the  nerves  extending  thence 
into  the  adjacent  structures,  representing  one  of  these  enlarged 
gastrula-territories.  In  point  of  fact,  however,  a  single  gastrula 
had  formed  the  basis  of  it  all,  the  structures  being  gradually 
elaborated  from  this  with  progress  in  development ;  but  it  will 
serve  the  purpose  of  illustration.  But  the  same  mechanical 
principle  applies  for  producing  afflux  and  efflux  of  the  fluids 
in  the  intestine  of  the  worms  as  in  the  primitive  intestine  of 
the  gastrula ;  only,  in  place  of  escaping  through  the  oral  orifice 
by  reflux  action  during  contraction,  it  is  passed  onward  into 
the  next  adjacent  gastrula  territory,  which  expands  simulta- 
neously for  reducing  pressure  in  order  to  compel  it  in  this 
direction  ;  otherwise,  it  would  escape  through  the  oral  orifice  by 
reflux  action  as  in  gastrula.  And  thus,  by  producing  areas  of 
low  and  high  pressures  contiguous  to  each  other,  a  dual  force 
applies  in  the  worms  for  increasing  the  action  in  the  fluids,  at 


Pig.  14. — Nervous  System  of  the  Leech  (Owen)  ;  a,  anterior  cerebral  ganglia  ;  b  b,  the 
10  ocelli  arranged  around  the  margin  of  the  upper  lip  ;  c,  posterior  cerebral  ganglia. 

the  same  time  compelling  them  to  move  onward  in  the  canal, 
and  flowing  always  from  high  to  low  pressure  in  conformity 
with  universal  law :  the  one  acting  as  a  pulling,  the  other  as 
a  pushing-force  upon  the  food,  acting  simultaneously.  This, 
then,  is  the  explanation  for  the  undulations  which  pass  along 
the  body  of  the  worms  during  ingestion,  indicating  the 
passage  of  the  food  along  the  canal  of  the  intestine,  as  before 
remarked.  Nothing  could  be  more  simple  or  more  effective. 
The  principle  is  not  new  either,  but  old — old  as  Nature ! 
Only  it  had  not  been  applied  to  the  animal  body  for  elucidating 
the  vital  phenomena ;  as  though  the  body  could  be  outside 
and  independent  of  this  fundamental  force  in  Nature  for  con- 
trolling the  movement  of  the  fluids  upon  which  the  very  body 
itself  is  based. 


60        COORDINATION   OF  INTERNAL   AND   EXTERNAL   PAUTS. 


One  other  circumstance,  of  great  importance  in  this  connec- 
tion, remains  for  mention,  namely  : 

The  Progressive  Increase  in  Size  with  Ingestion. — As  there 
is  no  unoccupied  space  in  the  body,  it  follows  there  must  be 
corresponding  increase  in  size  with  ingestion,  a  circumstance 
which  also  has  forcible  illustration  in  the  worms  (Figs.  15,  16, 
17);  the  gorged  leech,  for  example  (Fig.  17),  being  several 
times  the  natural  size  in  the  empty  condition  (Fig.  15).  This 
is  all  brought  about  gradually,  the  waves  of  expansion  pass- 
ing  from  the  mouth-cavity  along  the  body  of  the  sucking 


Pig.  15. 


Fig.  16. 


Fig.  17. 


Three  cute  of  the  Leech,  showing  the  appearance  before  imbibition,  during  imbibition, 
and  after  imbibition. 

Fig.  15. — Appearance  before  imbibition. 

Fig.  16. — Appearance  during  imbibition  ;  I,  oral  sucker,  expanded  ;  2,  wave  of  expan- 
sion passing  along  the  body  to  the  hinder  portions  ;  3,  where  they  are  added  up  in 
the  general  expansion  which  takes  place. 

Fig.  17. — Appearance  after  imbibition,  and  when  the  animal  is  fully  gorged  with  blood 
and  further  expansion  is  impossible. 

leech  (Fig.  16,  1,  2),  being  merged  in  the  general  expansion 
(3)  which  results.  In  other  words,  room  is  made  for  the  food 
as  it  progresses  in  the  body  ;  otherwise  the  balance  in  pressure 
could  not  be  maintained,  and  the  food  would  regurgitate.  And 
this  thing  continues  till  the  limit  in  expansion  is  reached,  when 
no  more  food  can  be  taken  (Fig.  17). 

And  but  for  the  fact  that  the  internal  and  external  parts  are 
in  correspondence,  how  otherwise  could  this  action  be  pro- 
duced %    Thus,  the  stomach,  which  extends  nearly  the  whole 


THE    PRINCIPLE  APPLIED   TO   RESPLEATION.  61 

length  of  the  body,  is  not  forcibly  pulled  open  by  the  action 
in  the  muscular  envelope,  but  expands  of  its  own  accord 
in  harmonious  concert  with  the  muscular  envelope  ;  while  for 
effecting  this  action  in  the  walls,  as  also  for  producing  the  local 
actions,  is  the  office  of  the  double  ganglionic  chain  of  nerves 
upon  the  ventral  surface,  with  the  nerves  extending  thence  into 
the  parts,  whereby  the  utmost  concert  of  action  is  produced, 
so  that  not  only  the  requisite  changes  in  pressure  for  swallow- 
ing the  food  may  be  readily  effected,  but  at  the  same  time  a. 
general  expansion  in  the  body  commensurate  with  the  food 
ingested,  for  maintaining  a  balance  in  pressure,  may  also  be 
produced — the  one  involving  the  other  ;  as  the  requisite  room- 
must  be  made  in  the  body  as  fast  as  the  food  is  taken  in, 
in  order  to  maintain  a  balance  in  pressure  ;  otherwise  impos- 
sible. 

All  of  which  is  plain  enough  and  easily  understood.  Of 
course,  as  the  food  is  removed  by  the  absorptive  processes, 
the  intestine  contracts  correspondingly,  while  the  fullness  in 
the  vessels  which  this  produces  effects  corresponding  expan- 
sion in  tliem,  for  which  a  special  physiological  adjustment 
obtains  in  the  organism,  the  whole  being  regulated  by  nervous 
force,  to  be  spoken  of  further  on.  We  are  now  prepared  to 
take  another  important  step  in  the  physiological  adjustments, 
notably : 

The  Action  in  the  Lungs. — This  necessity  for  maintaining' 
correspondence  between  the  internal  and  external  surfaces,  in 
order  to  produce  afflux  and  efflux  of  the  fluids  in  the  body- 
interior,  must  now  be  applied  to  the  mechanics  for  producing 
afflux  and  efflux  of  air  and  blood  in  the  lungs  for  respiratory 
purposes,  the  principle  being  the  same  In  a  word,  the  lungs 
and  containing  walls  expand  and  contract  simultaneously,  as  be- 
fore remarked,  and  which  is  not  only  true  logically,  but  it  also 
admits  of  actual  demonstration. 

For  this  purpose  selection  is  made  of  the  frog,  which  has 
neither  diaphragm  nor  ribs  for  assisting  the  action  in  the 
lungs,  the  ribs  remaining  in  an  undeveloped  and  rudi- 
mentary condition  (Fig.  18)  ;  and  the  lungs  being  contained 
in  the  common  visceral  cavity,  offer  exceptional  advantages 
for  studying  respiration,    which  is  thus,    so   to   speak,    ex- 


G2 


THE    PRINCIPLE  APPLIED   TO   RESPIRATION. 


posed  and  laid  bare.  It  is  the  common  impression  among 
physiologists  and  naturalists  that  the  frog  respires  by  pump- 
ing the  air  in  and  out  of  the  lungs  by  means  of  the  throat-ap- 
paratus, but  which  is  only  partially  true,  the  arrangement  in 
the  throat  serving  to  assist  it  simply  ;  and  that  it  is  not  the 
fundamental  circumstance  in  batrachian  respiration,  1  have 
fully  proven  by  excising  a  portion  of  the  floor  of  the  mouth,  so 
as  to  lay  the  cavity  open  and  effectually  destroy  it  as  a 
pumping  apparatus  for  pumpirig  air  into  the  lungs  :  notwith- 
standing this,  however,  the  animal  continued  to  respire,  ulti- 
mately recovering  from  the  wound,  which  closed  by  cicatriza- 
tion in  the  course  of  two  weeks.     The  immediate  effect  of  the 


Fig.  18.— Skeleton  of  the  Frog.— Owen. 

operation  was  very  characteristic,  the  body  suddenly  col- 
lapsiug  like  a  pricked  balloon  over  the  whole  lung-region 
(Fig.  19),  the  spine  and  the  stumps  of  the  undeveloped  ribs 
standing  out  in  great  prominence,  produced  by  the  soft  tissues 
upon  the  sides  being  forcibly  compressed  under  them,  impart- 
ing an  excavated  appearance  to  the  lateral  regions  ;  while  the 
abdominal  portions  (2)  appeared  round,  full  and  pendulous 
from  compression  of  the  envelope  around  them,  making 
them  also  stand  out  prominent.  The  collapse  was  so  sud- 
den and  unexpected  as  to  startle  me.  Fully  expecting 
him  to  die  outright,  which  I  thought  a  matter  of  course, 
and  saying  to  myself,  "I  am  not  going  to  prove  anything 
by  you,"  I  dropped  him  into  a  waste-bucket  containing  a  thin 


THE    PRINCIPLE  APPLIED   TO   RESPIRATION. 


63 


stratum  of  water  and  some  loose,  wet  paper;  and  the  night 
being  far  advanced,  I  at  once  sought  my  couch,  deeming 
the  experiment  over  and  done  with.  The  following  morn- 
ing, however,  upon  glancing  into  the  bucket,  the  animal,  to 
my  unspeakable  amazement,  was  sitting  up,  fully  expanded, 
head  high,  and  looking  at  me  out  of  his  great  luminous  eyes, 
as  natural  to  all  appearances  as  ever,  save  the  wound  in  his 
throat  (Fig.  20).  That  thin  stratum  of  water  in  the  bucket, 
together  with  the  loose  wet  paper,  under  which  he  could 
creep  to  keep  himself  moist,  were  the  best  possible  conditions 
for  promoting  recovery  after  the  operation,  though  I  was  not 
conscious  of  this  at  the  time  when  I  threw  him  into  it  with  a 


Fig.  19. 


Fig.  30. 


Two  cuts  of  the  Frog,  showing  the  collapsed  condition  of  the  body  produced  by  destruc- 
tion of  the  throat-apparatus,  and  the  subsequent  expansion  which  takes  place. 
Fig.  19. — Appearance  after  destruction  of  the  throat-apparatus  ;  1,  1,  outline  of  the 

undeveloped  ribs  ;  3,  abdominal  viscera  ;  3,  opening  in  the  floor  of  the  mouth. 
Fig.  20. — The  Frog  expanded  to  its  normal  size,  notwithstanding  the  opening  in  the  floor 

of  the  mouth. 

feeling  of  sore  disappointment  at  my  heart  over  the  untoward 
result  of  an  experiment  from  which  I  had  expected  so  much, 
and,  as  I  thought,  reasoned  out  so  carefully.  And,  after  all, 
there  he  was,  fully  alive,  and  on  the  qui  vive,  too,  as  though 
expecting  me  But  other  surprises  were  in  store  for  me,  if 
possible  of  a  still  more  agreeable  nature.  Thus,  I  found  while 
the  throat-apparatus  continued  to  move  to  and  fro,  the  wound 
alternately  widely  gaping  and  contracting,  but  still  leaving  a 
considerable  opening,  at  the  rate  of  120  to  130  per  minute,  that 
another  much  larger  and  slower  movement  was  pervading  the 
body  itself,  from  20  to  30  times  per  minute.     And  that  this 


64  THE    PRINCIPLE   APPLIED   TO   RESPIRATION. 

was  the  true  respiratory  movement  was  proven  by  the  fact 
that  it  corresponded  with  the  action  in  the  alee  nasi,  which  ex- 
panded and  contracted  synchronously,  as  is  the  case  in 
impeded  respiration,  and  which  at  once  stamps  its  true 
character.  The  destruction  of  the  throat-apparatus  and  the 
embarrassment  to  respiration  which  this  produces,  brings  into 
prominence  the  action  in  the  lungs  and  containing  walls,  espe- 
cially in  the  lateral  dorsal  regions,  where  it  is  conspicuous. 
In  times  of  excitement,  both  movements  are  increased,  but  the 
relative  frequency  is  maintained;  showing  that  they  are  also 
connected. 

Upon  taking  him  up,  however,  he  struggled  violently  in  my 
hands  and  again  collapsed,  presenting  in  all  respects  the  same 
appearances  as  at  first.  But  replacing  him  at  once  in  the 
bucket,  he  filled  himself  out  again  with  air  as  before.  Of 
course,  the  action  is  necessarily  comparatively  slow,  from  the 
abseuce  of  "  ribs"  and  special  muscles  for  assisting  the  lungs 
by  the  leverage  they  afford  for  reducing  pressure ;  still,  he 
accomplishes  it,  nevertheless,  filling  the  lungs  completely  and 
restoring  the  natural  rotundity  (Fig.  20).  Moreover,  he 
seemed  to  acquire  more  power  in  the  larynx  for  holding  the 
air,  not  collapsing  so  readily  as  at  first,  an  amount  of  compen- 
sation for  the  injured  throat-apparatus  taking  place  in  this  way. 

Eight  days  after  the  operation,  in  exhibiting  him  to  Dr. 
G unth er,  the  distinguished  naturalist  of  the  British  Museum, 
and  others,  and  holding  him  firmly  by  the  hind  legs  for  the 
purpose,  by  a  sudden  and  energetic  movement  leaped  out  of 
my  hands,  and  falling  with  great  impact  against  the  floor,  again 
collapsed  as  previously.  It  produced  a  sensation.  The  wound 
had  contracted  considerably,  but  the  top  of  the  larynx  was 
easily  visible  through  the  opening.  A  week  subsequently, 
however,  it  had  closed  entirely,  and  the  old  condition  was  com- 
pletely restored.  But  there  was  some  deformity  in  the  throat 
from  loss  of  substance  and  contraction  of  the  wound,  bring- 
ing the  contour  of  the  larynx  into  prominence,  and  showing 
something  like  an  Adam's  apple.  But  he  was  thoroughly 
recovered,  hale  and  hearty,  and  swam  with  greatest  eager- 
ness in  the  tank  when  placed  with  the  other  frogs  at  the  shop 
where  I  obtained  him. 


THE    PRINCIPLE   APPLIED   TO   RESPIRATION.  05 

It  is  scarcely  necessary  to  add  that  the  results  of  this  opera- 
tion prove  incontrovertibly  that  the  lungs  expand  synchro- 
nously with  the  containing  walls  ;  otherwise  it  were  utterly  im- 
possible for  the  animal  to  have  inflated  them  or  filled  them 
with  air,  having  neither  diaphragm  nor  "ribs,"  nor  a  throat- 
apparatus  for  compelling  air  into  the  lungs.  And  since  air 
flows  from  high  to  low  pressure,  it  follows  that  the  lungs  must 
expand  in  order  to  produce  afflux  of  air  in  the  organs  ;  other- 
wise impossible.  But,  then,  might  not  expansion  in  the 
muscular  envelope  itself  effect  it  ?  No  ;  this  should  produce 
low  pressure  in  the  vi3ceral  cavity,  and  thereby  speedily 
bringing  life  to  an  end  by  causing  afflux  of  the  fluids  in  the 
cavity,  and  arresting  the  functions  in  the  lungs.  In  short, 
the  low  pressure  must  be  made  within  the  lungs  themselves  in 
order  to  produce  afflux  and  efflux  of  air  and  blood  in  the 
organs  for  respiratory  purposes,  at  the  same  time  keeping 
them  in  their  respective  channels  ;  otherwise  impossible  At 
any  rate,  however,  the  muscular  envelope  expands  ;  and  if  this 
expands,  there  is  no  reason  on  earth  why  the  lungs  may  not 
do  likewise — nay,  must  do  so  in  the  very  nature  of  things,  the 
law  compelling  this  circumstance. 

Explanation  for  the  Sudden  Collapse  of  the  Body-  Walls. — 
In  batrachians  the  tongue  is  applied  to  the  posterior  nares  in 
the  form  of  a  valve,  which  converts  the  mouth-cavity  into  an 
air-tight  chamber  and  enables  the  pumping  movements  to  be 
made  in  the  throat  for  assisting  respiration,  at  the  same  time 
it  prevents  escape  of  the  air  when  rude  force  is  applied,  as 
when  the  animal  leaps ;  the  absence  of  ' '  ribs' '  for  imparting 
solidity  and  firmness  to  the  walls  rendering  this  necessary. 
Hence,  when  the  air  chamber  is  destroyed,  the  animal  strug- 
gling, collapse  is  inevitable,  the  larynx  itself  not  being  sufficient 
for  the  purpose.  It  subserves  other  important  uses  also. 
Thus  it  enables  the  animal  to  remain  for  considerable  periods 
under  the  water,  by  pumping  the  air  to  and  fro,  or  backward 
and  forward,  between  the  chamber  and  the  lungs,  till  fully 
exhausted  of  its  oxygen,  when  it  returns  to  the  surface  for 
more  ;  the  air  being,  so  to  speak,  bottled  up  during  this  time. 

Chelonia  and  crocodilia  all  possess  this  power  ;  so  also  ceta- 
cea  ;  the  enormous  chambers  forming  the  tortuous  nasal  pass- 


06    MODE   OF   COORDINATING   THE   LUNGS   AND   BODY-WALLS. 

ages  in  the  latter,  which  are  closed  externally  by  means  of  a 
plug  projected  from  the  side,  subserving  this  purpose  and 
enabling  the  animals  to  disappear  for  long  periods  beneath 
the  surface 

In  the  seal  the  external  nares  are  closed  by  powerful  muscles, 
which  are  promptly  contracted  at  the  moment  of  disappear- 
ance. Other  animals  are  similarly  endowed.  Being  air- 
breathers,  the  air  must  be  thus  forcibly  retained  during  the 
time  of  submergence  ;  otherwise  they  could  not  live. 

Mode  of  Effecting  Coordination. — The  mode  the  lungs 
are  coordinated  with  the  muscular  envelope  or  containing 
walls,  together  with  the  changes  and  mode  of  distribution 
of  the  nerves  to  the  parts,  is  also  easily  explained,  in  the 
worms,  for  example,  we  have  seen  that  coordination  of  the  in- 
ternal and  external  parts  forming  the  enteron  and  muscular 
envelope  or  containing  walls,  is  effected  by  means  of  a  double 
chain  of  nervous  ganglia  extending  along  the  ventral  surface, 
whence  the  nerves,  which  are  given  off  laterally,  also  symmet- 
rically (Fig.  14)  proceed  directly  into  the  adjacent  structures, 
passing  from  one  to  the  other,  the  intimate  relations  they  sus- 
tain to  each  other  enabling  this  to  be  very  readily  done  ;  but 
with  progress  in  development  and  the  differentiation  of  a 
visceral  cavity,  however,  which  is  necessary  for  increasing  the 
action  in  the  viscera  in  connection  with  the  special  functions, 
it  is  manifest  that  this  calls  for  fresh  arrangement  in  the 
nerves  for  coordinating  the  viscera  with  the  muscular  envelope 
and  containing  walls,  since  they  are  now  completely  separated. 
Hence,  the  pneumogastric  nerves  which  are  sent  down  from 
above  for  connecting  the  viscera  with  the  medulla  oblongata 
(Fig.  21)  ;*  and  being  correlated  with  the  spinal  nerves  to 
the  external  parts,  concert  of  action  is  readily  produced,  so 
that  the  whole  performs  as  but  a  single  organ  only  for  pump- 
ing aii*  and  blood  through  the  alveoli  for  respiratory  purposes. 
And  in  the  later  stages  in  development,  when  a  diaphragm 
comes  into  the  scheme  for  separating  the  viscera  in  the  abdo- 
men from  the  viscera  in  the  chest,  likewise  the  phrenic  nerves 
(Fig.  91,  F)  for  coordinating  this  portion  with  the  external 
parts,  so  that  the  utmost  harmony  is  produced  throughout, 

*  For  the  purpose  of  illustration  simply  being  fundamentally  the  same  as  in 
the  frog. 


MODE   OF   COORDINATING  THE  LUNGS   AND   BODY-WALLS      67 

the  correlation  of  all  these  nerves  in  the  medulla  oblongata 
enabling  this  to  be  very  readily  done.  There  is  increasing 
complexity,  but  the  principle  in  the  mechanics  furnishes  a 
key  for  easily  unraveling  the  structures,  which  are  means 
to  ends  simply.  The  existence  of  the  pneumogastric  nerves, 
then,  is  readily  accounted  for.     The  role  they  perform  in  the 


Fig.  31. — Left  Pneumogastric  Nerve,  in  diagram  (Dalian).  1,  pharyngeal  branch  ; 
2,  superior  laryngeal ;  3,  inferior  or  recurrent  laryngeal  nerve  ;  4,  pulmonary 
branches  ;  5,  6,  stomach,  liver,  etc. 

viscera  in  the  abdomen  will  come  up  later,  but  its  importance 
would  be  difficult  to  overestimate.  In  the  fishes  we  have 
the  analogue  of  this  in  the  opercula  and  branchia,  together 
with  the  branches  of  the  8th  pair  (Fig.  166)  for  setting  up  the 
pumping  actions  for  producing  afflux  and  efflux  of  the  water 


08        PORTAL   CIRCULATION   INCREASED   BY   RESPIRATION. 

and  blood  in  these  organs  i'or  r  ■  spiratory  purposes,  while  the 
other  portions  are  continued  on  into  the  abdomen  and  the 
external  muscular  walls  (Fig.  15S,  o,  4)  for  coordinating  them 
with  the  branchia  and  opercula,  or  the  same  as  in  the  air- 
breather,  the  principle  being  the  same.  And  so  likewise 
down  the  whole  chain  in  development  to  where  a  visceral 
cavity  ceases  to  exist,  similar  arrangements  obtain  for  co- 
ordinating the  viscera  with  the  muscular  envelope  appropriate 
to  the  stage  in  development  and  the  requirements  in  respira- 
tion and  circulation,  in  order  to  evolve  the  force  expended  in 
them.  But  for  the  sake  of  brevity,  we  pass  over  the  details, 
the  matter  being  so  very  obvious. 

Dependence  of  the  Portal  Circulation  upon  Respiration. — 
Finally,  we  have  to  mention  a  circumstance  of  deepest  import 
in  the  mechanics  of  circulation,  namely  :  The  rhythmical  com- 
pression of  the  viscera  in  the  abdomen  synchronous  with 
respiration,  and  which  is  produced  by  respiration,  for  increas- 
ing the  portal  circulation  correspondingly,  and  which  it 
would  be  difficult  to  overestimate,  since  the  portal  circula- 
tion could  not  be  carried  on  without  it ;  neither  could  a  bal- 
ance be  maintained  in  the  absorptive  and  oxygenating  proc- 
esses which  the  scheme  calls  for,  and  which  requires  the 
absorptive  processes  in  the  intestine  and  the  portal  circulation, 
to  be  connected  with  respiration  in  such  manner  that  the  effort 
to  respire  must  necessarily  increase  the  two  correspondingly. 
It  is  easily  apprehended.  Thus,  the  muscular  envelope  form- 
ing the  walls  of  the  common  visceral  cavity  functions  as  the 
Jloor  of  support  to  the  viscera  in  the  abdomen,  which  are 
everywhere  in  contact  with  them,  gravitation  compelling 
this  circumstance  (Figs.  19,  2 ;  25,  U),  hence  are  necessarily 
affected  by  every  movement  in  the  walls  for  effecting  the 
changes  of  pressure  in  the  lungs  for  producing  afflux  and 
efflux  of  air  and  blood  in  the  organs  for  respiratory  purposes, 
and  which,  of  course,  should  increase  the  portal  circulation 
correspondingly.  It  could  not  do  otherwise,  in  the  very 
nature  of  things.  They  are  forcibly  compressed  during  ex- 
piration for  producing  high  pressure  in  the  lungs,  being  com- 
pressed against  them  for  this  purpose,  when  the  heart  serves 
as  a  suction-force  upon  the  portal   blood,  which  thus  flows 


PORTAL   CIRCULATION   INCREASED   BY   RESPIRATION.         69 

from  high  to  low  pressure ;  while  during  inspiration  the 
suction-force  in  the  lungs  is  added  to  this,  which  greatly 
increases  the  action  by  means  of  which  the  portal  circula 
tion  is  readily  carried  on  ;  otherwise  impossible.  For  show- 
ing the  dependence  of  the  portal  circulation  upon  res- 
piration, the  following  physiological  experiment  possesses 
fresh  interest,  and  is  certainly  very  conclusive  upon  this 
matter :  * 

' '  Two  frogs  are  slightly  curarized  and  placed  side  by  side  in 
the  supine  position.  In  both,  the  heart  and  great  vessels  are 
exposed,  as  in  the  preceding  section.  It  having  been  ascer- 
tained that  the  circulation  is  normal  in  each  animal,  and  the 
frequency  of  the  contractions  having  been  noted,  the  brain 
and  spinal  cord  are  destroyed  in  one  of  the  frogs,  by  inserting 
a  strong  needle  into  the  spinal  canal  immediately  below  the 
occipital  bone,  and  then  passing  it  upward  and  downward. 
This  may  usually  be  accomplished  without  much  loss  of  blood. 
If  now  the  frog  which  has  been  deprived  of  its  nervous  centres 
is  compared  with  the  other,  it  is  seen  that  in  the  former, 
although  the  heart  is  beating  with  perfect  regularity  and  un- 
altered frequency,  it  is  empty,  and  in  consequence,  instead 
of  projecting  from  the  opening  in  the  anterior  wall  of  the 
chest,  it  is  withdrawn  upward  and  backward  toward  the 
oesophagus.  The  emptiness  of  the  heart  is  not  limited  to  the 
ventricle  and  bulb.  The  auricles  are  alike  deprived  of  blood ; 
and  if  the  heart  is  drawn  forward  by  the  apex,  it  is  seen  that 
the  sinus  wnosus  and  vena  cava  inferior  are  in  the  same  con- 
dition. The  state  is  therefore  not  dependent  upon  any  cause 
inherent  in  itself  but  on  the  fact  that  no  blood  is  conveyed  to 
it  by  the  veins.  To  make  this  still  more  evident,  the  rest  of 
the  visceral  cavity  may  be  opened,  when  it  is  seen  that 
although  the  vena  cava  is  collapsed,  the  intestinal  veins  are 
distended." 

The  distinguished  physiologist  and  experimentalist,  in  cast- 
ing about  for  an  explanation  of  the  preceding  phenomena,  con- 
ceived the  idea  that  the  non-arrival  of  the  blood  in  the  heart 
was  due  to  relaxation  or  expansion  of  the  arterial  system  from 
vaso-motor  paralysis  ;  and  while  this  undoubtedly  was  a  factor 

*  Hand-book  for  the  Physiological  Laboratory. — Burdon-Sartderson. 


70        POKTAL   CIRCULATION   INCREASED   BY   RESPIRATION. 

in  the  case,  it  would  certainly  not  account  for  a  portion  of  the 
phenomena — notably,  the  turgescence  in  the  portal  vessels. 
If  loss  of  tension  in  the  arterial  system  produced  emptiness 
in  the  lower  cava-system  then,  how  could  it  produce  the  oppo- 
site condition  of  turgescence  in  the  adjacent  portal  system  % 
If  true  in  the  one,  it  should  be  true  in  the  other  also.  We  take 
it,  however,  that  suspension  of  respiration  was  the  principal 
cause  of  the  phenomena,  for  the  reason  that  it  arrested  the 
pumping  action  in  the  lungs  for  aspirating  the  systemic  capil- 
laries, and  the  rhythmical  compression  of  the  viscera  in  the 
abdomen,  which  this  produces  for  increasing  the  portal  circu- 
lation. 

The  extent  and  firmness  of  the  liver-structure,  the  character 
of  the  portal  vessels,  and  the  firm  adhesion  of  the  walls  of  the 
hepatic  veins  to  the  liver- substance  for  maintaining  patency, 
all  tend  to  show  this  circumstance.  And  since  there  is  no  other 
means  for  increasing  circulation  but  by  rhythmical  changes  in 
pressure,  it  follows  that  the  liver  must  be  rhythmically  com- 
pressed for  increasing  circulation  in  it.  Moreover,  it  should  be 
borne  in  mind  that  the  portal  vessels  are  sandwiched,  so  to 
speak,  between  two  extensive  capillary  systems  :  one  in  the 
intestines,  in  relation  with  the  mucosa  ;  the  other  in  the  liver- 
substance.  Hence  the  necessity  for  this  direct  application  of 
force  for  increasing  circulation  in  the  liver  and  portal  vessels. 

The  flow  of  the  portal  blood  into  the  heart  when  the  two  are 
connected  by  a  canula,  serves  to  establish  two  important 
facts  :  1.  That  the  portal  circulation  is  obstructed  in  the  liver 
when  respiration  is  suspended,  the  blood  in  consequence 
rapidly  accumulating  in  this  system  of  vessels.  2.  That  the 
action  in  the  heart  is  inadequate  for  the  portal  circulation, 
which  requires  a  greater  force  than  this  for  effecting  it  In 
short,  the  body  being  based  upon  pressure  and  the  power  of 
producing  rapid  rhythmical  changes  in  pressure,  nothing  short 
of  the  pumping  action  in  the  abdomen,  which  is  set  up  by 
respiration,  and  the  rhythmical  changes  in  pressure  w  hich  this 
produces  in  the  viscera,  would  be  sufficient  for  the  purpose. 
And  when  we  get  a  little  further  on  in  the  mechanics,  it  will 
•be  shown  how  this  action  in  the  abdomen  for  increasing  the 
portal  circulation  is  mnde  to  connect  with  the  action  in  the  in- 


DEPENDENCE   OF   POETAL   AND   SYSTEMIC    CIECULATICN.     71 

testines  for  increasing  the  digestive  and  absorptive  processes 
for  producing  correspondence  and  maintaining  a  balance  in 
this  circulation,  inclusive  of  the  action  in  the  lymph- 
channels,  which  cannot  be  omitted  in  the  scheme.  The  rapid 
absorption  of  fat,  albumen,  alcohol,  which  are  non-dialyzable, 
hence  will  not  pass  the  membranes  without  mechanical  force, 
are  alike  included  in  it.  In  fine,  the  explanation  of  the  portal 
circulation,  to  be  satisfactory,  should  embrace  the  entire  me- 
chanics for  increasing  circulation  in  the  abdomen,  and  for 
maintaining  this  in  correspondence  with  the  physiological 
requirements  in  the  organism  ;  and  which,  of  course,  should 
include  the  extensive  nervous  connections  subsisting  in  the 
viscera,  together  with  the  arrangements  of  the  structures  in 
the  organs  as  means  to  ends  All  of  which  have  ample  explana- 
tion. It  comes  to  this,  then,  that  the  heart  is  not  able  to 
carry  on  the  portal  circulation,  as  is  fully  proven  by  the  above 
experiment. 

But  that  is  not  all  that  is  proven  by  this  experiment.  ISay  ! 
not  by  half  and  more.  Put  this  great  fact  in  the  foreground 
of  the  mental  picture :  When  the  respiratory  centre  is  destroyed, 
both  the  systemic  andportal  circulations  arepromptly  arrested; 
this  notwithstanding  the  heart  is  beating  with  perfect  regular- 
ity and  unaltered  frequency  !  The  mechanical  work  this  organ 
is  capable  of  is  far  too  inadequate  for  the  enormous  labor 
which  is  involved  in  the  circulation,  the  non-arrival  of  the 
blood  at  the  heart,  the  empty  condition  of  the  chambers,  the 
"collapsed"  vense  cavee,  together  with  the  swollen  and  turgid 
portal  system,  are  all  eloquent  of  that  fact ;  showing  conclu- 
sively that  the  cardinal  circumstance  in  the  circulation  is  the 
great  pumping  movements  which  oire  set  up  by  respiration, 
extending  from  the  lungs  to  the  tissue-territories  and  from 
surface  to  surface  of  the  body ;  for  to  the  movements  takings 
place  in  the  chest  and  abdomen  there  are  rhythmical  expansions 
and  contractions  of  the  arterial  and  capillary  systems,  inclusive 
of  the  venous,  synchronous  with  respiration  for  producing  cor- 
respondence and  maintaining  a  balance  in  the  circulation ; 
otherwise  impossible. 

But  with  progress  in  development  ' '  ribs' '  are  differentiated 
in  the  external  walls,  or  muscular  envelope,  with  a  system  of 


72  RESPIRATION   IN   BIRDS. 

"internal"  and  "external  intercostal  muscles' '  for  operating 
tli em  in  connection  with  the  action  in  the  lungs,  the  other 
muscles  in  the  envelope  also  powerfully  assisting,  whereby  the 
action  in  the  lungs  is  greatly  increased.  This  arrangement  sub- 
serves a  double  purpose  :  it  protects  the  lungs  against  the  rude 
experiences  to  which  the  animal  is  exposed,  so  as  to  prevent 
"  collapse'"  of  the  organs  ;  at  the  same  time  it  enables  the  ex- 
pansions and  contractions  to  be  more  readily  effected.  In 
short,  it  energizes  the  action  in  the  lungs.  But  as  the  action  in 
the  "ribs"  is  exhaustively  treated  in  existing  works,  it  will  not 
be  necessary  to  consider  it  here  further  than  to  refer  to  the  action 
the  diaphragm  exerts  upon  them,  which  will  be  taken  up  after 
briefly  referring  to  respiration  in  birds,  the  matter  having 
escaped  attention. 

Respiration  in  Birds. — Passing,  for  the  sake  of  brevity, 
over  the  intervening  stages  in  development,  as  the  action  is 
fundamentally  the  same  throughout,  and  for  the  reason,  also, 
that  it  is  so  conspicuous  in  them,  moreover  so  abundant  and 
accessible,  we  take  up  respiration  in  the  birds ;  and  being, 
also,  warm-blooded,  this  brings  us  nearer  to  the  higher  animals, 
where  interest  culminates.  The  structures  in  birds  are  homolo- 
gous with  the  reptilian  and  mammalian,  upon  either  side  of 
them,  the  special  modifications  that  obtain  in  them  being 
adaptive  changes  to  the  stage  in  development  simply.  But  as 
yet  no  diaphragm  appears  for  separating  the  viscera  in  the 
abdomen  from  the  viscera  in  the  chest,  which  comes  into  the 
scheme  in  the  succeeding  stage  in  development  as  a  special 
adjustment  to  the  altered  mechanics  in  the  intestinal  canal 
for  increasing  the  digestive  and  absorptive  processes,  the 
portal  and  lymph  circulations  which  will  be  taken  up  after 
the  systemic  circulation  shall  have  been  disposed  of  as  they 
can  be  treated  more  advantageously  by  following  this  method  ; 
besides,  that  method  should  be  adopted  in  the  treatment 
which  would  connect  all  the  phenomena,  anatomical  and 
physiological,  in  one  harmonious  whole,  and  make  everything 
plain  and  easily  understood.  The  point  we  wish  to  empha- 
size in  this  connection,  however,  is,  that,  notwithstanding  the 
absence  of  a  diaphragm,  the  birds  respire  easily  and  readily, 
can  produce  high  vocal  resonance,  carry  whole  notes,  and  make 


RESPIRATION   IN   THE  ABSENCE   OF   A   DIAPHRAGM.  73 

all  the  varied  sounds  peculiar  to  them,  showing  that 
respiration  is  not  only  under  voluntary  control,  and  all 
the  parts  fully  coordinated,  but  at  the  same  time  is  very 
energetic.  And  as  development  is  by  addition  and  differ- 
entiation, in  order  to  understand  respiration  in  mamma- 
lia, respiration  in  birds  and  reptiles  would  first  have  to 
be  understood.  The  question,  then,  resolves  itself  into  this  : 
What  is  the  mechanics  for  producing  the  rhythmical  changes 
in  pressure  within  the  lungs  of  birds  for  compelling  afflux  and 
efflux  of  air  and  blood  in  the  organs  for  respiratory  purposes  ? 
— when  it  will  be  in  order  to  inquire  as  to  the  role  in  the  dia- 
phragm for  still  further  increasing  the  action.  Briefly,  respira- 
tion in  birds  is  accomplished  by  means  of  rhythmical  expan- 
sions and  contractions  in  the  abdomen  and  chest,  producing  a 
to-and-fro  movement  of  the  viscera  in  the  longitudinal  axis  of 
the  body  ;  the  viscera  in  the  abdomen  being  in  contact  with  the 
lungs,  fitting  accurately  against  them  and  moving  in  and  out  of 
the  excavation,  piston-like,  under  the  action  of  the  muscles  in  the 
abdomen  ;  while  the  lungs  are  in  concert  with  this  movement, 
expanding  and  contracting  simultaneously  and  synchronously, 
whereby  low  and  high  pressures  are  alternately  produced 
within  the  lungs  themselves,  which  the  scheme  calls  for  for 
producing  afflux  and  efflux  of  air  and  blood  in  the  alveoli,  at 
the  same  time  confining  the  fluids  within  their  channels ; 
otherwise  impossible.  This,  then,  is  the  explanation  of  the 
pumping  action  in  the  abdomen  of  the  birds  ;  the  expansile 
action  corresponding  with  inspiration,  the  contractile  with  ex- 
piration. And  when  at  rest  upon  the  ground,  the  hinder  parts 
are  raised  and  lowered  alternately  by  this  action  in  the  abdo- 
men, and  is  so  conspicuous  that  the  respirations  may  be  readily 
counted  at  a  distance  from  the  body  ;  rising  during  inspira- 
tion and  falling  again  during  expiration.  Brief  reference  to 
the  anatomy  is  necessary. 

Descriptive  Anatomy. — In  the  first  place,  the  costal  frame- 
work forms  an  ovoidal  cavity  ;  the  small  contracted  end  pre- 
senting anteriorly  at  the  root  of  the  neck,  while  the  large 
expanded  end,  presenting  posteriorly,  is  open  and  in  free  com- 
munication with  the  abdomen,  some  of  the  viscera  lying  with- 
in it  (Fig.  22).     The  skeleton  itself,  however  (Fig.   23),   gives 


74 


RESPIRATION   IN   THE   ABSENCE    OF   A   DIAPHRAGM. 


no  idea  of  the  extensive  abdominal  cavity  which  forms  the 
large  end  of  the  common  ovoidal  chamber.  But  by  contrast- 
ing the  cuts  a  mental  picture  may  be  formed.  For  example, 
the  long,  narrow  innominata  (Fig.  23,  s,  s' ,  s"),  one  for  each 
side,  together  with  the  narrow  os  sacrum,  form  the  roof  of 
the  cavity  ;  while  the  £>osterior  (p)  and  lateral  walls  are  formed 
of  soft  parts,  principally  muscles,  which  are  homologous  with 
those  in  the  higher  animals.     But  in  order  to  fully  expose  the 


Fig.  22. — Longitudinal  sect'on  of  the  Thoracic- Abdominal  Cavity  iu  the  Hen,  showing 
the  viscera  in  situ.  The  outline  of  the  ovoidal  cavity  is  unpaired  by  allowing  the 
acetabulum  (k)  to  remain.  A,  ventriculus  callosus  ;  B,  liver  ;  c,  intestines  ;  d,  kid- 
ney ;  c,  lungs  ;  F,  heart  ;  h,  left  pulmonary  artery  ;  g,  left  anonymous  artery ; 
i,  ovisacs  ;  k,  acetabulum  ;  /,  glenoid  cavity  ;  m,  ribs  ;  n,  process  of  sternum  ;  o, 
pectoral  muscles  ;  p,  abdominal  walls ;  r,  nasal  orifices ;  S,  proventriculus ;  s, 
oesophagus  ;  L,  atmosphere. 

whole  of  the  abdominal  portion  of  the  chamber,  however,  the 
whole  forming  a  perfect  ovoid,  the  innominatum  also  would  have 
to  be  removed,  thus  bringing  into  view  the  whole  of  the  intes- 
tines, a  few  coils  only  now  presenting  (Fig.  22,  c),  the  rest 
being  contained  in  the  pelvic  excavation.  The  anterior  ex- 
tremity of  the  bone  corresponds  with  d,  the  posterior  with  the 


RESPIRATION   IN  THE  ABSENCE   OF   A   DIAPHRAGM.  75 


Fig.  23.— Skeleton  of  a  Fowl.— Chauveau.  Frcm  A  to  B,  cervical  vertebras— 1,  spinous 
process  of  the  third  vertebra  ;  2,  inferior  ridge  on  body  of  the  same  ;  3,  styloid  pro- 
longation of  the  transverse  process  of  the  same  ;  4,  vertebral  foramen  of  the  same  : 
1',  2',  3',  4',  the  same  parts  in  the  twelfth  vertebra.  From  B  to  C,  dorsal  vertebrae  : 
6,  spinous  process  of  the  first  ;  7,  crest  formed  by  the  union  of  the  other  spinous  pro- 
cesses. From  D  to  E,  coccygeal  vertebrae  ;  F,  G,  head  ;  8,  interorbital  septum  ;  9, 
foramen  of  communication  between  the  two  orbits  ;  10,  prem axillary  bone  ;  10', 
external  openings  of  the  nose  ;  11,  maxilla  ;  12,  square  bone  ;  13,  jugal  bone  ;  H, 
sternum  ;  14,  brisket,  or  keel ;  15,  episternal  process  ;  16,  internal  lateral  process ; 
17,  lateral  external  process  ;  18,  membrane  which  closes  the  internal  notch  ;  19, 
membrane  of  the  external  notch  ;  J,  etc.,  superior  ribs  ;  20,  posterior  process  of  the 
fifth  ;  J,  inferior  ribs  ;  K,  scapula  ;  L,  coracoid  bone  :  M,  furculum  ;  m,  m,  its  two 
branches  ;  N,  humerus  ;  O,  ulna  ;  o,  radius  ;  P,  P7,  bones  of  carpus  ;  Q,  Q',  bones 
of  metacarpus  ;  B,  first  phalanx  of  the  large  digit  of  the  wing  ;  r,  second  phalanx  of 
the  same  ;  B',  phalanx  of  thumb  ;  S,  ilium  ;  S',  ischium  ;  £",  pubis  ;  21,  sciatic 
foramen  ;  22,  foramen  ovale  ;  T,  femur  ;  U,  patella  ;  V,  tibia,  X,  fibula  ;  F,  single 
bone  of  tarsus  ;  Y,  metatarsus  ;  23,  superior  process,  representing  a  united  metatarsal 
bone  ;  24,  process  supporting  the  claw  ;  Z,etc,  digits. 


76  ADJUSTMENTS    IN   THE   VISCERA. 

commencement  of  the  soft  abdominal  walls,  midway  between 
p  and  A\ 

By  contrasting  the  corresponding  parts  in  the  skeleton 
(Fig.  23)  some  idea  may  be  formed  of  the  length  and  depth 
of  the  pelvic  excavation,  and  the  great  extent  of  the  muscular 
structures  forming  the  sides  and  floor  of  the  abdomen.  Now, 
then,  in  regard  to  the  mechanics  in  respiration.  As  will  be 
seen,  the  solid  and  heavy  viscera,  notably  the  ventriculus  callo- 
sus  and  liver,  are  at  the  very  bottom  of  the  cavity,  the  liver 
resting  against  the  projecting  sternal  processes  (Figs.  22,  A,  B; 
23,  H,  16),  with  all  the  other  viscera  resting  upon  them  as  their 
floor  of  support,  the  ventriculus  itself  resting  entirely  upon 
the  soft  abdominal  walls  (Fig.  22,  A,p).  The  sternum,  by 
projecting  long,  slender  processes  toward  the  abdomen,  aids 
considerably  in  supporting  the  liver,  but  the  weight  in  the 
greater  portion  of  the  viscera  is  sustained  by  the  abdominal 
muscles  simply,  and  these,  by  being  inserted  in  the  sternal  pro- 
cesses, aid  in  supporting  the  liver  also,  together  with  the  overly- 
ing viscera,  so  that  all  are  necessarily  affected  by  the  action  in 
the  walls.  Finally,  the  stomach  (ventriculus)  and  liver  are 
firmly  secured  to  the  posterior  walls  by  means  of  duplicatures 
of  the  lining  membrane  (Fig.  22,  A,  B,  p),  the  lobes  of  the  liver 
extending  far  down  the  sides  of  the  ventriculus  in  form  of 
a  saddle,  and  secured  by  means  of  connective  tissue  and  the 
overlying  peritoneum,  so  that  the  two  are  compelled  to  move 
together  under  the  action  in  the  walls.  It  results  from  this 
arrangement  in  the  parts,  that  with  every  expansion  in  the 
abdomen  the  viscera  are  pulled  downward  and  backward,  gravi- 
tation and  the  traction  from  the  retaining  ligaments  compel- 
ling this  circumstance  :  while  during  contraction  they  are 
pushed  in  the  opposite  direction,  or  forward  and  upward.  In 
the  former,  they  tend  to  pull  away  from  the  lungs,  thereby 
tending  to  produce  low  pressure  in  the  visceral  cavity  itself  ; 
but  as  this  would  defeat  the  end  in  view,  the  lungs  expand 
simultaneously  and  par I  passu  with  expansion  in  the  abdomen 
for  maintaining  the  intra-abdominal  pressure,  keeping  in  close 
apposition  with  the  viscera  for  this  purpose ;  which  has  the 
effect  of  confining  the  low-pressure  area  within  the  lungs 
themselves  for  aspirating  the  air  and  blood  for  respiratory 


MODE  OF  PKODUCLTSTG  AFFLUX  AND  EFFLUX  IN  THE  LUNGS.  77 

purposes,  otherwise  impossible,  since  the  fluids  flow  from  high 
to  low  pressure,  in  conformity  with  universal  law.  During 
contraction,  however,  when  the  action  is  reversed,  the  viscera 
are  firmly  compressed  against  the  lungs,  which  maintains 
intra-abdominal  pressure,  at  the  same  time  that  high  intra- 
pulmonic  pressure  which  this  produces  causes  the  fluids  to  flow 
out  of  the  organs  again  till  pressure  is  uniform,  the  lungs  also 
contracting  pari  passu  with  contraction  in  the  abdominal 
muscles  for  increasing  the  action.  Finally,  for  effecting  co- 
ordination we  have  the  pneumogastric  and  spinal  nerves,  or  the 
same  as  in  the  frog,  the  principle  being  the  same.  So,  then, 
at  no  time  is  there  any  difficulty  in  producing  the  requisite 
changes  in  pressure  in  the  alveoli  for  compelling  afflux  and 
efflux  of  air  and  blood  in  the  chambers  for  respiratory  pur- 
poses, at  once  the  object  and  purpose  of  all  these  arrangements. 
But  did  not  the  lungs  expand  during  inspiration  in  the  manner 
alleged,  the  effect  would  be  to  develop  low  pressure  wit  kin 
the  abdomen  itself,  which  would  defeat  the  end  in  view  by 
causing  afflux  of  the  fluids  in  this  locality,  in  consequence 
rapidly  putting  an  end  to  life.  Not  outside  of,  then,  but 
within  the  lungs  themselves  must  low  pressure  be  made  dur- 
ing inspiration,  in  order  to  keep  the  fluids  in  their  relative 
channels,  as  before  remarked. 

This,  then,  is  the  explanation  for  the  pumping  action  so  con- 
spicuous in  the  abdomen  of  the  birds ;  and  having  no  dia- 
phragm for  pulling  the  viscera  away  from  the  lungs  during 
inspiration,  they  are  fastened  by  means  of  duplicatures  of  the 
lining  membrane  to  the  posterior  abdominal  walls,  thereby 
obviating  the  necessity  for  a  diaphragm.  And  with  the  big 
end  of  the  excavation,  containing  the  greater  portion  of  the 
viscera,  the  heavier  at  the  bottom,  the  whole  inclined  at  an 
angle  of  about  30  degrees  (Fig.  22),*  gravitation  would  neces- 
sarily favor  inspiration,  enabling  expansion  to  be  more  readily 
made  by  adding  its  force  to  the  force  in  the  muscles,  thereby  im- 
parting energy  to  the  movement.  The  result  of  this  action  is  to 
compel  the  air  (L)  to  flow  into  the  air-passages  and  alveoli 
through  the  nasal  orifices  (r)  till  pressure  is  uniform,  the  ve- 

*  In  the  skeleton,  the  student  must  look  from  the  abdomen  upward,  filling 
out  the  soft  tissues  in  his  mind,  in  order  to  get  the  angle  of  inclination. 


78  MODE  OF  PRODUCING  AFFLUX  AND  EFFLUX  IN  THE  LUNGS. 

nous  blood  at  the  same  time  flowing  into  the  alveolar  plexuses 
from  every  portion  of  the  body,  the  heart  serving  to  assist  the 
action,  as  has  already  been  stated  :*  while  for  compelling 
them  out  again,  we  have  simply  to  reverse  the  action  in  the 
abdomen  and  lungs  in  the  manner  as  stated — the  one  flowing 
out  by  reflux  action  through  the  route  of  ingress,  the  other 
into  the  left  chambers  of  the  heart  and  arterial  system  on  its 
way  to  the  cell-brood,  the  heart,  of  course,  assisting  the  ac- 
tion ;  otherwise  it  were  impossible  to  produce  the  high  press- 
ure in  the  arterial  system  for  increasing  the  capillary  circula- 
tion commensurate  with  the  physiological  requirements,  in 
order  to  generate  the  force  which  is  expended,  the  main- 
tenance of  the  upright  position  itself  alone  demanding  rapid 
circulation  for  effecting  it,  and  which,  of  course,  would 
involve  rapid  respiration,  as  in  no  other  way  could  a  balance  be 
maintained.  Hence  the  rapid  pumping  action  in  the  »abdomen 
of  the  birds.  But  while  this  action  is  going  on  in  the  abdomen 
and  lungs,  however,  the  chest  itself  expands  and  contracts 
simultaneously  for  increasing  the  action,  thereby  enabling  the 
lungs  to  expand  outward  as  well  as  downward,  which  the 
action  in  the  lungs  calls  for.  But  it  is  not  so  extensive  as 
the  action  in  the  abdomen,  save  in  the  portions  contiguous 
to  the  abdomen,  into  which  the  muscles  of  the  latter  have 
insertion,  notably  the  long,  slender  sternal  processes  (Fig.  23, 
J3,  16,  17)  and  adjacent  ribs,  and  which,  of  course,  are  com- 
pelled to  respond  to  the  action  in  the  muscles,  thus  making 
the  movement  in  respiration  conspicuous  in  them  ;  the  length 
of  the  muscles  in  the  abdomen  being  the  explanation  of  this 
circumstance,  since  the  amount  of  elongation  and  shortening 
in  the  muscles  is  determined  by  their  length ;  hence,  the 
rhythmical  expansions  and  contractions  in  the  abdomen  dur- 
ing respiration  are  necessarily  much  greater  than  in  the  chest. 
Morever,  the  ribs  in  birds  do  not  partially  revolve  around 
the  long  axis  for  flaring  them  open  upon  the  sides,  as  in  other 
animals,  the  long  costal  processes  (20)  effectually  preventing 
it,  this,  together  with  the  forked  articulations  to  the  spine,  for 

*  The  author  indulges  the  hope  that  the  importance  and  gravity  of  the  sub- 
ject will  serve  as  apology  for  frequent  reiteration  of  the  cardinal  points  in 
order  to  enforce  attention  and  stamp  them  indeliby  upon  the  mind. 


MODE   OF  INCREASING   IT  IN   FLYING.  79 

imparting  firmness  to  this  portion  of  the  framework,  as  the  base 
of  support  to  the  wings.     At  the  same  time,  however,  we  have 
to  note  a  compensatory  arrangement  for  maintaining  expansile 
and  contractile  action  in  the  chest ;  and  this  consists  in  a  series 
of  cartilaginous  articulations  in  the  mid-costal  region,  dividing 
the  ribs  into  two  portions,  or  "superior"  and  "inferior"  ribs 
(Fig.  23,  I,  J),  which  join  each  other  at  an  obtuse  angle,  form- 
ing an  elbow  ;  while  below  they  connect  with  the  sternum  by 
terminal  cartilages,  in  the  usual  way;  so  that  the  chest  can 
readily  be  expanded  or  contracted  by  the  overlying  muscles, 
being  forcibly  compressed  by  means  of  the  contracted  abdomi- 
nal muscles   during  expiration,   and  springing  forcibly  out 
again  with  expansion  in  the  muscles  during  inspiration  ;  the 
pectoral  muscles  also  assisting  in  the  action,  especially  during 
flight.     And  as  this  arrangement  in  the  ribs  Occurs  only  in 
birds,  it  is  highly  probable  this  is  the  case  ;  so  as  to  compel 
respiration  to  be  in  correspondence  with  the  increase  in  the 
activities.     Thus,  the  powerful  pectoral  muscles  covering  the 
whole  anterior  and  lateral  portions  of  the  chest,  have  their 
broad  points  of  origin  in  the  keel  (Fig.  23,  14)  (the  widely- 
expanded  process  projected  from  the  central  portions  of  the 
sternum  for  this  purpose),  while  the  points  of  insertion  are  in 
the  head  and  proximal  portions  of  the  shaft  of  the  humerus  ; 
hence,  every  effort  to  extend  and  retract  the  wings  would  tend 
to  expand  and  contract  the  chest  correspondingly,  the  points 
of  origin  and  insertion  being  movable ;  consequently,  would 
have  to  be  separated  or  approximated  by  the  expansions  and 
contractions  in  the  muscles,  as  this  would  increase  or  diminish 
their  length  correspondingly.   In  this  manner,  then,  respiration 
is  increased  with  the  flight,  which  is  the  physiological  finality 
in  this  adjustment ;  at  the  same  time,  however,  it  enables  the 
energetic  action  in  the  lungs,  for  producing  loud  vocal  reso- 
nance, to  be  more  readily  made.     In  this  condition  the  pos- 
terior portions  of  the  chest  are  forcibly  compressed  by  the 
energetic  contraction  of  the  abdominal  muscles,  thereby  dimin- 
ishing the  transverse  with  the  longitudinal  axis  in  the  visceral 
cavity,  and  which,  of  course,  must  effect  corresponding  com- 
pression of  the  lungs  for  producing  the  high  intra-pulmonic 
pressure  which  is  essential  for  producing  the  vibrations  in  the 


80  CIRCULATION   IN   THE  AIR-SACS   AND   BONES. 

vocal  cords ;  the  one  involving  the  other.  And  in  order  to 
obtain  some  idea  of  the  enormous  role  performed  by  the  viscera 
and  the  muscles  in  the  abdomen  in  respiration,  one  has  need 
only  to  watch  the  action  of  these  parts  in  the  screaming  parrot, 
and  he  will  perceive  that  the  viscera  are  more  and  more  com- 
pressed into  the  chest-excavation  by  the  action  in  the  mus- 
cles as  the  notes  are  prolonged  or  raised  in  volume,  the  abdomen 
in  consequence  becoming  more  and  more  flat  and  reduced  in 
size,  till  at  the  end  of  the  note  the  greater  portion  of  the  viscera 
are  in  the  chest  excavation ;  at  the  same  time,  the  lower  chest- 
walls  are  firmly  held  down  by  the  muscles  in  the  abdomen,  that 
the  force  may  be  concentrated  in  the  lungs.  At  the  end  of  the 
effort  they  fly  back  to  the  original  positions  under  the  ener- 
getic expansion  in  the  muscles,  aided  by  the  resiliency  in  the 
ribs,  in  which  force  is  stored  during  contraction,  to  be  yielded 
up  again  during  the  subsequent  expansion,  so  that  nothing  is 
lost,  but  only  carried  over  from  one  rhythmic  movement  to 
the  other 

In  quiet  respiration,  however,  the  motion  in  the  trunk 
seems  limited  to  the  abdomen  and  the  proximal  portions  of 
the  chest.  In  addition  to  this  mode  of  respiration  in  the  lungs, 
the  birds  are  provided  with  air-sacs,  the  whole  thoracic- 
abdominal  cavity  being  divided  by  septa  of  serous  membrane 
into  numerous  inter-communicating  chambers,  all  of  which 
are  freely  supplied  with  air,  brought  into  them  through  the 
bronchial  tubes,  which  open  upon  the  lung  surface,  passing 
through  and  through  the  lung  substance  for  this  purpose 
(Fig.  24,  a,  e),  the  air  being  admitted  to  the  very  bones  them- 
selves ;  and  in  birds  of  powerful  flight  penetrating  even  be- 
tween the  muscles  of  the  neck  and  limbs.  And  it  is  easy  to  per- 
ceive how  the  action  in  the  abdomen  and  chest  should  increase 
this  circulation  also  by  expanding  and  contracting  the  air- 
sacs  correspondingly,  at  the  same  time  compelling  the  air  into 
and  out  of  the  bones  with  afflux  and  efflux  of  air  in  the  lungs, 
and  enabling  them  to  readily  inflate  the  sacs  for  buoying  them 
in  the  media  when  swimming  or  flying,  by  diminishing  the 
body-density,  and  increasing  surface  correspondingly.  It  is 
needless  to  extend  the  matter. 

In  conclusion  :  It  is  manifest,  from  this  relation  of  the  parts 


PORTAL   CIECULATIOlSr. 


81 


in  the  abdomen,  that  the  pumping  action  in  respiration  should 
increase  the  portal  circulation  correspondingly.  Thus,  the 
liver  substance  being  spread  out,  as  it  were,  against  the  firm 


Pig.  34.— Lungs,  Heart,  and  Great  Vessels  of  Buteo  Vulgaris.—  Gegenbaur.  tr, 
trachea  ;  i,  crop  ;  ae,  communication  between  the  air-sacs  and  the  lungs  ;  6,  bursa 
f  abricii  ;  ao,  aortic  arch  :  aad,  art.  anonyma  destra  ;  aas,  art.  anonyma  sinistra  ; 
ps,  art.  pulmonalis  sinistra  ;  c,  carotid  ;  am,  visceral  artery ;  vci,  commencement 
of  the  inferior  vena  cava  :  vcm,  vena  coccygeo-mesenterica. 


82  PORTAL   CIRCULATION. 

ventriculus  (Fig  22,  B,  A),  with  the  other  viscera  resting  upon 
them,  must  necessarily  undergo  a  degree  of  compression  with 
every  expiration,  and  increasing  till  the  end  of  this,  which 
would  tend  to  force  the  blood  through  the  open  hepatic  veins, 
the  aspiratory  force  in  the  heart,  which  is  close  to  the  organ,  being 
added  to  this  ;  while  during  inspiration  the  organ,  by  spring- 
ing back  again  to  the  original  shape,  should  produce  a  suction- 
force  upon  the  portal  blood,  at  the  same  time  the  suction  force 
in  the  lungs  is  added  to  that  in  the  heart  for  aspirating 
the  blood  in  the  liver;  so  that  inspiration  must  necessarily 
increase  circulation  in  the  liver.  It  must  do  so,  in  the  very 
nature  of  things.  Of  course,  the  same  applies  for  the  viscera 
in  the  abdomen,  connected  with  the  lower  eava  system 


CHAPTER  V. 

RESPIRATION  IN   MAMMALIA. 

Respiration  in  the  Mammalia  Fundamentally  the  Same  as  in  Birds  and  Reptiles,  the  Lungs 
and  the  Muscles  in  the  Abdomen  Expanding  and  Contracting  Simultaneously — Office 
of  the  Diaphragm  and  the  Occasion  for  it— The  Action  in  the  Diaphragm  Alternating 
with  the  Muscles  in  the  Abdomen,  the  One  Contracting  as  the  Other  is  Expanding,  and 
Vice  Versa — The  Action  the  Diaphragm  Exerts  Upon  the  Ribs,  Bending  and  Flaring 
Them  Open  Upon  the  Sides  During  Inspiration,  and  Vice  Versa  During  Expiration — 
Mode  of  Coordination,  Inclusive  of  the  Lungs,  the  Whole  Moving  in  Perfect  Con- 
cert— Circumstances  in  the  Vital  Phenomena,  Anatomical  and  Physiological,  which 
Make  it  Absolutely  Certain  that  the  Lungs  Expand  and  Contract  Regularly  and 
Rhythmically  Synchronous  with  the  Actions  in  the  Diaphragm  and  Containing 
Walls — Physiological  Experiments  Proving  This  Circumstance — The  Action  in  the 
Tracheal  System — Mode  of  Maintaining  Cleanliness  in  the  Alveoli  and  Air-Pass- 
ages— Significance  of  a  Cough — The  Action  of  the  Trachea  in  Vocalization — Expla- 
nation for  the  Devious  Course  of  the  Recurrent  Laryngeal  Nerves,  which  First 
Descend  Into  the  Chest  to  Connect  with  the  Lungs  Before  Proceeding  to  the  Larynx 
and  Vocal  Cords,  Ascending  Thence  Upon  the  Trachea  for  This  Purpose,  No  Matter 
How  Long  the  Neck  May  Be  ;  e.  g. ,  the  Giraffe. 

This  brings  ns  to  Respiration  in  the  Mammalia.  From 
what  has  preceded,  it  follows  that  the  first  thing  to  engage 
the  attention  of  the  student  of  physiology  is  the  relation  which 
the  viscera  sustain  to  the  muscular  envelope  or  containing 
walls,  since  it  is  by  means  of  the  rhythmical  expansions  and 
contractions  taking  place  in  the  latter  that  the  relative  changes 
in  pressure  are  produced  in  the  former  for  compelling  afflux  and 
efflux  of  air  and  blood  in  the  aveoli  for  respiratory  purposes  ; 
the  viscera,  of  course,  being  responsive  to  the  movements  in 
the  walls  in  order  to  effect  these  results,  and  in  close  contact 
with  them  all  the  while.  In  short,  that  the  principal  force  for 
producing  the  requisite  changes  in  pressure  for  effecting  respi- 
ration in  mammals  inheres  in  the  muscular  envelope  itself, 
with  which  the  internal  parts  are  coordinated  by  means  of  the 
nervous  apparatus,  as  obtains  in  the  preceding  stages  in 
development,  the  principle  being  the  same ;  only  there  is 
increasing  differentiation  in  the  organs,  with  the  development 


84  FLOOR   OF   SUPPORT   TO   THE   VISCERA. 

of  a  special  organ  or  structure  (diaphragm)  for  separating  the 
viscera  in  the  abdomen  from  the  viscera  in  the  chest,  and 
which  is  rendered  necessary  by  the  changes  which  have 
occurred  in  the  intestines  for  increasing  the  digestive  and 
absorptive  processes  commensurate  with  the  force  which  is 
expended  in  these  animals,  which  are  womb-bearers,  and  oc- 
cupy a  higher  plane  in  development.  In  present  physiology 
the  weight  in  the  viscera  receives  little  attention,  which  is  a 
great  oversight,  as  the  whole  of  them,  with  the  entire  body 
itself,  have  adjustment  with  gravitation,  so  that  it  would  be 
utterly  impossible  to  explain  the  phenomena  in  the  absence  of 
this  essential  factor.  iN  ow  that  attention  is  called  to  it,  the 
necessity  for  this  strong  floor  of  support  to  the  viscera  in  the 
abdomen  will  at  once  appear  obvious,  since  the  folds  of  lining 
membrane  forming  the  ligaments  for  retaining  them  in  their 
relative  positions,  at  the  same  time  permitting  free  movement 
in  them  in  connection  with  the  special  functions,  contain  the 
nerves  and  blood  vessels  to  the  viscera,  which  would  at  once 
inhibit  much  traction  upon  them  ;  otherwise  inevitable  but 
for  this  floor  of  support.  The  weight,  too,  is  very  consid- 
erable, approximating  one-fifth  of  the  weight  in  the  body. 

So,  then,  we  can  readily  understand  why  the  heavy  viscera 
should  occupy  the  bottom  of  the  cavity,  while  the  floor  of  sup- 
port should  be  substantial,  the  liver  itself  resting  against  the 
sternum,  with  the  stomach  and  intestines  against  the  muscular 
walls.  But  in  order  to  fully  appreciate  the  mechanics,  it  will  be 
necessary  to  again  refer  to  the  special  anatomy  in  the  parts. 

Descriptive  Anatomy.— As  will  be  seen,  the  muscular  en- 
velope still  functions  as  the  floor  of  support  to  the  viscera  (Fig. 
25,  i£),  though  the  relative  positions  they  sustain  to  each  other 
are  somewhat  changed.  Thus,  while  the  liver  (L)  is  still  in 
front  of  the  stomach  (S),  and  resting  against  the  sternum  and 
ribs,  as  in  the  birds  (Fig.  22,  B),  it  is  not  intimately  attached 
to  the  stomach,  but  is  attached  to  the  incurvated  surface  of  the 
diaphragm  instead  by  duplicatures  of  the  lining  membrane; 
while  the  stomach  itself,  now  widely  expanded,  is  in  easy  con- 
tact with  it  simply,  and  free  to  move  in  the  visceral  cavity,  that 
it  may  function  both  as  a  receptacle  for  the  food  and  at  the 
same  time  effect  digestive  action  in  it ;  and  that  the  intestines, 


FLOOR   OF   SUPPORT   TO   THE   VISCERA. 


85 


also  greatly  increased  in  size,  are  not  now  resting  upon  the 
liver  and  stomach  as  the  floor  of  support  (Fig.  22,  c,  c,  B,  A), 
but  occupy  a  position  posteriorly  and  resting  against  the  mus- 
cular floor  itself,  in  easy  contact  with  them  (Fig.  25,  i",  C — E, 
S,  L) ;  at  the  same  time,  are  also  free  to  move  within  the 
cavity  in  connection  with  respiration  and  the  special  functions 
in  the  organs  ;  while  they  are  retained  in  their  relative  positions 
by  means  of  duplicatures  of  the  lining  membrane,  forming  the 
gastric,  meso-colic  and  mesenteric  ligaments.  From  this  rela- 
tion of  the  parts,  it  follows  that  the  viscera  are  compelled  to 


Fig.  25. — Diagrammatic  Longitudinal  Section  of  the  Abdomen  in  the  Horse,  showing  the 
position  of  the  Viscera,  and  the  relations  they  sustain  to  the  lower  abdominal  walls, 
or  floor  of  support.  E,  abdominal  walls  ;  S,  stomach  ;  L,  liver  ;  O,  omentum  ;  C, 
colon  ;  M,  mesentery  ;  I,  small  intestines  ;  Q,  duodenum  ;  P,  pancreas  ;  B,  bladder  ; 
V,  vagina  ;  B,  rectum  ;  d,  diaphragm  ;  k,  oesophagus  ;  A,  aorta. 

respond  to  every  movement  in  the  muscular  walls.  It  could 
not  be  otherwise,  in  the  very  nature  of  things.  Thus,  during 
inspiration,  when  the  muscles  expand,  the  viscera  sink  down, 
so  to  speak,  with  the  walls  ;  but  when  contraction  sets  in  dur- 
ing expiration,  they  rise  again  with  them,  in  this  manner  effect- 
ing a  to-and-fro  movement  of  the  viscera  in  the  chest-excavation 
during  respiration  for  producing  the  rhythmical  changes  of 
pressure  in  the  lungs  ;  while  the  large  apron-like  duplicatures 
of  the  lining  membrane  forming  the  great  omentum  (o)  inter- 
vening between  them  and  the  floor,  facilitates  the  gliding  move- 
ments and  obviates  friction,  the  secretions  in  the  lining  membrane 
serving  also  to  lubricate  the  organs,  thus  reducing  friction  to  a 
minimum,  the  mechanics  being  fundamentally  the  same  as  in 


86  OFFICE   OF   THE   DIAPHRAGM. 

the  birds  and  reptiles.  Now,  then,  in  regard  to  the  diaphragm. 
In  addition  to  the  great  increase  in  size  of  the  stomach  and 
intestines,  they  are  now  filled  and  distended  with  air  (Fig.  25, 
8,  C,  31),  which  tends  to  balloon  them  in  the  cavity,  and  but 
for  this  diaphragm  for  restraining  and  operating  them  in  con- 
nection with  respiration,  would  rise  up  in  the  chest-excavation 
and  inhibit  the  action  in  the  lungs,  putting  an  end  to  life. 
Hence  this  circumstance. 

At  the  same  time,  however,  this  calls  for  the  phrenic  nerves 
(Fig.  91,  F),  which  are  correlated  with  the  other  nerves  in  the 
medulla  oblongata  for  operating  the  organ  in  connection  with 
the  muscular  envelope  and  lungs,  all  the  parts  acting  in  har- 
monious concert  as  previously,  in  order  to  produce  the 
changes  of  pressure  in  the  alveoli.  In  fine,  the  old  founda- 
tions are  simply  built  upon  and  further  extended  for  produc- 
ing a  larger  amount  of  work  commensurate  with  the  force 
expended  in  the  organism,  while  .  the  principle  in  the  me- 
chanics remains  unchanged  and  unchangeable,  from  the  very 
nature  of  things.  Thus,  with  inspiration  the  diaphragm  con- 
tracts for  pulling  the  viscera  away  from  the  lung-region,  while 
the  abdomen  expands  simultaneously  and  pari  passu  with 
this  action,  in  order  to  make  requisite  room  for  descent  of  the 
diaphragm  and  lungs,  which  expand  synchronously  with  the 
muscles  in  the  abdomen,  as  before  ;  while  during  expiration 
the  action  is  reversed  simply,  the  diaphragm  expanding 
simultaneously  and  pari  passu  with  contraction  in  the 
abdominal  muscles  and  the  lungs,  in  order  to  effect  this  action. 
In  short,  the  old  mechanics  continues  in  operation  as  it  ever 
did,  while  the  dia23hragm  comes  into  the  scheme  as  an  adapta- 
tion to  the  changes  effected  in  the  stomach  and  intestines, 
for  increasing  the  digestive  and  absorptive  processes,  and 
without  which  they  never  could  have  been  made,  leaving 
development  at  the  stage  in  the  birds.  And  let  it  not  be  ima- 
gined that  the  powerful  muscles  in  the  abdomen  are  made  to 
yield  to  the  force  in  the  diaphragm,  and  are  pulled  into  exten- 
sion by  this  means,  which  cannot  be  thought  of  for  a  single 
moment  even,  the  vast  preponderance  of  the  muscles,  of  itself 
alone,  inhibiting  this  action  ;  if,  forsooth,  such  rude  mechanics 
were  at  all  admissible. 


OFFICE   OF   THE   DIAPHRAGM.  87 

The  large  undulation,  extending  over  the  abdomen  during 
inspiration,  is  produced  by  expansion  in  the  muscles,  together 
with  contraction  in  the  diaphragm  for  pushing  the  viscera 
along  pari  passu  with  this  action  for  occupying  the  space  thus 
made,  in  this  manner  creating  the  billow  ;  while  during  ex- 
piration the  action  is  reversed,  the  contraction  which  then  sets 
in  in  the  muscles  of  the  abdomen  causing  the  wave  of  reflux 
in  the  viscera,  which  return  again  into  the  chest,  the  dia- 
phragm at  the  same  time  expanding  pari  passu  with  contrac- 
tion in  the  walls,  till  at  the  end  of  a  forced  expiration  most  of 
the  viscera  are  in  the  chest-excavation,  leaving  the  abdomen 
less  than  one-half  the  size  it  had  at  the  end  of  inspiration. 
Indeed,  in  an  athlete  this  recession  of  the  viscera  in  the  chest 
may  be  made  so  complete  by  the  energetic  action  of  the 
muscles  as  to  remove  nearly  all  the  lower  coils  of  the 
mesentery,  so  as  to  bring  the  walls  in  actual  contact  with  the 
lumbar  vertebrae  (Fig.  128),  which  are  easily  felt. 

Of  course,  the  diaphragm,  with  the  lower  chest-region,  ex- 
pands correspondingly  in  order  to  effect  this  action  ;  otherwise 
impossible.*  So,  then,  it  is  very  readily  perceived  that  the 
diaphragm  is  in  perfect  concert  with  the  muscles  in  the  abdo- 
men ;  only  that  the  action  is  reversed  in  them,  the  former  con- 
tracting as  the  latter  are  expanding,  and  vice  versa.  The 
extensive  aciion  which  this  involves  to  the  viscera  is  amply 
provided  for  by  the  long  ligament  in  the  mesentery  (Fig.  25, 
m),  which  permits  the  loose  coils  of  intestine  to  readily  glide 
over  each  other  for  the  purpose  ;  at  the  same  time,  they  are 
freely  lubricated  by  the  secretions  in  the  membrane. 

But  as  we  shall  have  occasion  to  refer  to  this  again,  in  con- 
nection with  the  functions  in  the  abdomen,  it  need  not  detain 
us  here,  and  we  pass  on  to  the  special  phenomena  in  respira- 
tion, taking  them  up  as  we  go  along. 

The  Action  the  Diaphragm  Exerts  upon  the  "Hibs." — 
The  relations  which  the  diaphragm  sustains  to  the  ribs,  and 
the  manner  they  are  affected  by  its  action,  are  in  keeping 
with  the   comprehensiveness   in   animal  mechanics   and   the 

*  But  as  life  advances  the  cartilages  become  more  and  more  ossified,  while 
fat  accumulates  in  the  viscera  ;  hence,  this  extreme  action  becomes  more  and 
more  difficult. 


88  DIAPHRAGM    OPERATING  THE   RIBS. 

wonderful  utilitarian  methods  which  everywhere  obtain, 
Thus,  while  contraction  in  the  diaphragm  increases  the  longi- 
tudinal axis  in  the  chest,  it  at  the  same  time  compels  the 
ribs  to  flare  open  upon  tlie  sides  for  increasing  the  trans- 
verse axis  simultaneously,  so  that  the  lungs  may  expand 
outward  as  well  as  downward  for  increasing  the  action  in 
them  correspondingly.  And  while  it  is  true  that  the  ribs 
expand  in  the  absence  of  a  diaphragm  for  effecting  it,  as  is 
fully  proven  in  birds  and  reptiles,  nevertheless  the  converse  is 
equally  true,  as  I  have  fully  demonstrated  upon  the  dog  and 
cat  by  denuding  the  chest  of  its  muscles  *  (the  animal  under 
chloroform)  without  embarrassing  respiration  in  the  least, 
while  the  ribs  continued  to  expand  and  contract  in  the  usual 
way.  In  point  of  fact,  the  intercostal  muscles  share  in  the 
general  muscular  paresis  which  accompanies  chloroform  nar- 
cosis, f  Moreover,  "  sex"  would  seem  to  exert  no  influence  in 
this  respect,  the  action  in  the  chest  and  abdomen  being  as  con- 
spicuous in  the  female  as  in  the  male,  there  seeming  to  be  no 
difference  in  them.  And  by  thus  eliminating  the  intercostal 
muscles,  chloroform  furnishes  a  crucial  test  of  this  action  in 
the  diaphragm  and  ribs.  The  explanation  of  the  mechanics  is 
sufficiently  easy,  notably :  The  diaphragm  is  attached  to  the 
ends  of  the  ribs,  being  inserted  in  the  cartilages  (Fig.  26,  9), 
whence  it  is  ballooned  in  the  chest  by  the  viscera  (Fig.  27)  so 
as  to  bring  the  muscular  fibres  in  correspondence  with  the 
long  axis  in  the  ribs  (A,f,f) ;  hence,  when  contraction  sets 
in  during  inspiration,  the  traction  would  necessarily  be  upon 
the  ends  of  the  costal  bows,  tending  to  bend  and  flare  them 
open  upon  the  sides  like  the  ribs  in  an  umbrella,  while  their 
resiliency  would  impart  elastic  action  to  expiration  with  relief 
in  the  traction  force  which  this  produces.  The  natural  curva- 
ture of  the  ribs,  being  also  slightly  twisted  upon  the  long  axis, 

*  In  destroying  the  intercostal  muscles,  great  care  is  necessary  to  prevent 
puncturing  the  pleurae,  which  is  extremely  difficult. 

•{•This  circumstance  I  had  seen  strikingly  illustrated  after  the  removal  of  a 
malignant  tumor  of  the  chest,  in  which  the  intercostals  were  laid  bare,  and 
which  flopped  to  and  fro  during  respiration,  being  pushed  in  and  out  alter- 
nately by  the  changes  in  pressure  within  the  chest,  which  continued  to  expand 
and  contract  in  the  usual  way. 


DIAPHRAGM   OPERATING  THE   RIBS. 


89 


Pig.  26. — Showing  the  manner  in  which  the  diaphragm  is  attached  to  the  cartilages  of 
the  inferior  ribs,  viewed  from  below. — Cbauveau.  1,1',  pillars  of  the  diaphragm  ; 
2,  left  pillar  ;  3,  common  tendon  ;  4,  tendinous  attachments  to  the  cartilages  ;  4', 
muscular  attachments  ;  5,  attachments  to  control  tendons  ;  6,  vena  cava  opening  ; 
7,  oesophagus  ;  8,  aorta  ;  10,  psoas  magnus  ;  11,  psoas  parvus  ;  12,  transverse  pro- 
cess of  lumbar  vertebra  ;  13,  superior  or  external  lumbar  muscles. 

Diaphragm  of  the  Horse  ;  Posterior  Face. — 1,  1',  the  two  portions  of  the  right  pillar  ;  2, 
left  pillar ;  3,  tendons  of  the  pillars  ;  4,  4',  peripheral  muscular  portion  ;  5,  left 
leaflet  of  the  aponeurotic  portion  ;  5,  5',  right  leaflet  of  the  same  ;  6,  posterior  vena 
cava  ;  7,  oesophagus  passing  through  the  opening  in  the  right  pillar  ;  8,  posterior 
aorta  between  the  two  pillars  ;  9,  cartilaginous  circle  of  the  ribs  ;  10,  11,  section  of 
the  psoas  muscle  ;  12,  section  of  a  lumbar  vertebra  ;  13,  section  of  the  common 
mass  ;  14,  retractor  muscle  of  the  last  rib  ;  15,  xiphoid  appendage  of  the  sternum. 


90  RESPIRATORY   ROCKING   IN   THE  BODY. 

must  inevitably  cause  them  to  revolve  in  the  long  axis  in  in- 
spiration, and  thus  flare  them  open  upon  the  sides. 

At  the  same  time,  the  external  intercostal  muscles  should  in- 
crease the  action  in  inspiration,  while  the  internal,  by  acting  in 
the  opposite  direction  during  expiration,  should  impart  corre- 
sponding energy  to  the  reverse  action,  so  that  the  force  in  these 
muscles  is  still  utilized  in  respiration,  though  not  essential  to 
it.  But  there  is  concert  of  action  throughout,  so  that  all 
act  together ;  and  which  is  readily  accomplished  by  the  cor- 
relation of  the  nerves  in  the  respiratory  centre,  the  intercostal 
nerves  supplying  the  external  parts,  inclusive  of  the  muscles  in 
the  abdomen  ;  the  phrenic  and  pneumogastric  nerves  the  in- 
ternal parts,  whereby  correspondence  is  produced  during 
inspiration  and  expiration  as  in  the  previous  stages  in  devel- 
opment. This  action  in  the  diaphragm  and  ribs  would  also 
explain  another  curious  circumstance,  notably  the  to-and-fro 
movement  in  the  head  and  shoulders  during  respiration,  or 
backward  and  forward,  otherwise  inexplicable.  1  n  other  words, 
the  head  and  shoulders  are  rocked  to  and  fro  by  the  action  in 
the  diaphragm  and  ribs  during  respiration,  and  the  deeper 
the  inspirations  the  more  conspicuous  it  becomes.  As  we 
have  said,  the  diaphragm  is  inserted  into  the  ends  of  the  ribs, 
and  with  the  viscera  against  its  breast  in  the  chest-excavation 
with  the  spinal  column  as  the  point  of  resistance,  it  is  mani- 
fest that,  with  contraction  in  the  diaphragm,  this  would  tend  to 
push  the  spine  backward  during  inspiration  ;  but  when  relief 
comes  during  expiration,  which  reverses  the  action,  the  spine 
would  return  to  its  original  position.  Hence  the  oscillations 
in  the  head  and  shoulders  during  respiration.  In  the  quad- 
ruped, this  circumstance  is  seen  in  the  sudden  shooting  for- 
ward of  the  whole  body,  produced  by  the  occasional  deep  and 
energetic  inspirations  which  the  animals  take,  especially  after 
rousing  from  slumber  and  during  digestion  ;  the  whole  body 
rocking  forward  upon  the  limbs  under  the  energetic  action  in 
the  diaphragm,  exerted  through  the  ribs  and  spinal  column. 
It  is  deeply  interesting.  One  other  circumstance  of  deep  im- 
port remains  for  mention  in  this  connection,  though  we  shall 
not  do  more  than  make  a  passing  reference,  as  the  matter 
pertains  more  particularly  to  another  work.*    Notably,  by 

*  See  "Gravitation  and  Development." 


EESPIKATORY   EOCKING  IN   THE  BODT. 


91 


the  action  in  the  diaphragm  the  viscera  are  thrown  against 
the  anterior  walls  of  the  abdomen  during  inspiration,  moving 
downward  and  backward  in  place  of  directly  backward  ;  the 
object  being  to  avoid  impact  in  the  pelvic  viscera,  which  would 
tend  to  force  out  the  contents  and  produce  strain  in  the  organs, 
hence  must  be  avoided.  And  by  attaching  the  diaphragm 
to  the  lumbar  vertebrae  and  the  ribs  in  the  manner  as  obtains, 


Fig.  27. — Section  of  the  chest-walls,  showing  position  of  the  diaphragm  in  the  excava- 
tion and  the  relations  it  sustains  to  the  lungs  and  ribs,  the  muscular  fibres  corre- 
sponding with  the  long  axis  in  the  ribs  (A,  F,  F).  A,  diaphragm  ;  B,  lungs  ;  C, 
heart ;  D,  pericardium,  reflected  ;  E,  sternum  ;  F,  ends  of  the  ribs. 

with  the  muscles  acting  in  the  long  axis  in  the  ribs,  this 
action  in  the  viscera  is  readily  effected,  so  that  impact  in  the 
pelvic  viscera  is  entirely  obviated,  which,  for  reasons  already 
given,  must  apply  for  every  stage  in  development  The  de- 
pression which  occurs  in  the  epigastric  region  during  inspira- 


92  ACTION    IN    THE   LUNGS. 

tion  is  due  to  the  bending  in  of  the  costal  and  ensif  orm  cartilages 
produced  by  the  traction  in  the  diaphragm  ;  but  as  life  ad- 
vances and  they  become  more  and  more  ossified,  it  is  less  and 
less  conspicuous  by  reason  of  the  rigidity  this  produces  in  the 
structures.  Thus,  these  adjustments  in  the  diaphragm  are  far- 
reaching. 

Concerning  the  Action  in  the  Lungs.— During  inspiration 
the  lungs  expand  as  the  diaphragm  contracts,  and  simulta- 
neously, expanding  with  a  force  sufficient  to  firmly  approxi- 
mate them  to  the  diaphragm  and  costal  surfaces,  for  obviating 
low  pressure  in  these  localities  and  producing  this  in  the 
alveoli  only,  expanding  outward  and  downward,  following  the 
diaphragm  and  completely  filling  the  pleural  cavities ;  while 
during  expiration  the  action  is  reversed,  the  organs  now  con- 
tracting and  moving  upward  and  inward,  the  ribs  against  the 
sides,  and  the  diaphragm  with  the  viscera  in  the  abdomen  push- 
ing against  the  bases  for  producing  high  pressure  in  the  alveoli, 
while  the  air  and  blood  flow  into  and  out  of  these  compart- 
ments simultaneously  in  order  to  equalize  pressure,  fluid  equi- 
librium compelling  them  to  do  so,  since  both  are  in  direct 
communication  with  these  localities,  each  by  means  of  a  special 
system  of  canals  in  correspondence  with  the  special' require- 
ments, the  veins  and  capillaries  being  also  very  soft  and  com- 
pressible. 

The  reasons  why  this  should  be  as  stated  are  simply  un- 
answerable, notably : 

1 .  Did  not  the  lungs  expand  pari  passu  with  contraction  in 
the  diaphragm,  the  result  would  be  to  develop  low  pressure  in 
the  pleural  cavities,  caused  by  the  pulling  away  of  the  dia- 
phragm and  ribs  from  the  lungs,  which  would  virtually  con- 
vert them  into  a  huge  cupping  apparatus  for  sucking  the 
fluids  into  the  pleural  cavities,  and  thereby  destroying  life  by 
producing  fatal  compression  of  the  lungs,  since  pressure  is 
transmitted  through  and  through  the  body  ;  and  the  fluids 
would  inevitably  flow  into  the  low-pressure  areas  to  equalize 
pressure.  It  could  not  be  otherwise,  in  the  very  nature  of 
things. 

Furthermore,  were  the  lungs  passive,  or  as  bags  simply, 
resistance  would  increase  with  the  traction  force  upon  them ; 


ACTION    IN    THE   LUNGS.  93 

hence,  during  inspiration,  there  should  be  progr ess ive  increase 
of  suction-force  in  the  pleurae,  caused  by  the  lungs  pulling 
one  way  and  the  diaphragm  with  the  chest-walls  the  other ; 
hence,  the  deeper  the  inspiration,  the  more  effective  it 
should  be  for  aspirating  the  fluids  into  the  cavities  of  'the 
pleurae. 

Tn  other  words,  a  force  which  acts  in  all  directions  and  upon 
every  fluid  without  distinction  cannot  be  made  subservient  to 
the  convenience  of  a  theory  which  would  limit  its  action  to  a 
single  fluid  (atmosphere),  and  confine  this  within  specific  limits 
or  to  the  lungs  simply,  and  ignoring  the  blood  altogether. 
Even  in  this  case,  however,  it  would  be  ineffectual,  since  the 
air  immediately  adjacent  to  the  pleurae  in  the  air-vesicles 
should  be  first  affected  before  the  outside  air  could  be  aspirated^ 
drawing  it  into  the  pleurae.  The  action  has  been  compared  to 
the  one  taking  place  in  a  "bellows;"  but  the  simile  fails 
at  an  essential  point,  for  there  never  was  a  bellows  with  a 
pair  of  lungs  inside  of  it  in  which  a  dual  circulation  of  air  and 
blood  is  carried  on.  And  though  the  law  of  pressure  which 
applies  is  the  same  in  both,  it  had  not  been  followed  to  its 
logical  results,  else  this  unfortunate  and  hasty  comparison,, 
tending  to  retard  physiological  inquiry,  would  never  have  been 
made. 

So,  then,  we  conclude  that  this  bellows-principle  cannot  pos- 
sibly be  the  mode  of  respiration,  but  that  the  lungs  perform  an 
active  role,  and  by  expanding  synchronously  with  expansion  in 
the  chest  following  the  descent  of  the  diaphragm  and  elevation 
of  the  ribs,  the  conditions  obtain  during  inspiration  for  produc- 
ing simultaneous  afflux  of  air  and  blood  in  the  alveoli  for  respi- 
ratory purposes,  at  once  the  object  and  purpose  of  the  mecha- 
nism ;  since  this  would  have  the  effect  of  producing  high  pres- 
sure in  the  pleurae,  with  low  pressure  in  the  alveoli,  making 
this  the  culminating  point  of  the  changes  in  pressure,  which  • 
the  scheme  calls  for.  While  during  expiration,  which  answers 
to  systole  in  the  lungs,  the  high  pressure  which  this  produces 
in  the  alveoli  causes  the  air  and  blood  to  flow  out  of  these  com- 
partments simultaneously,  the  former  by  reflux  action  through 
the  route  of  ingress,  the  latter  into  the  left  cardiac  chambers 
and  arterial  system  on  its  way  to  the  cell-brood  in  the  tissues, 


94  ACTION    IN    THE   LUNGS. 

the  objective  point  of  the  commerce,  as  before  remarked  ;  and 
the  whole  surface  of  the  organs  being  closely  embraced  by  the 
containing  walls  closing  tightly  around  them,  the  ribs  upon 
the  sides,  with  the  diaphragm  and  viscera  pressing  against 
their  bases  from  below,  must  necessarily  produce  high  pressure 
in  the  pleurae,  whereby  the  walls  of  the  alveoli  are  well  sup- 
ported for  obviating  rupture  during  expiration.  Hence,  at  no 
time  is  there  low  pressure  in  the  pleurae,  the  principle  under- 
lying the  mechanics  inhibiting  it,  as  we  very  well  know.  In 
this  manner,  then,  a  balance  is  maintained  for  keeping  the 
fluids  in  their  channels,  as  the  body  is  expanding  and  contract- 
ing regularly  and  rhythmically,  like  the  diastoles  and  systoles 
in  the  heart,  for  pumping  air  and  blood  through  the  alveoli 
simultaneously  for  respiratory  purposes,  while  the  demands  of 
imperious  law  are  fully  satisfied. 

So  much  for  the  principle  underlying  the  mechanics  in  res- 
piration, which  is  plain  enough 

2.  Under  this  head  are  included  a  number  of  reasons  apper- 
taining to  the  special  vital  phenomena,  anatomical  and  physio- 
logical, otherwise  inexplicable,  to  which  brief  reference  will 
be  made.  Indeed,  no  theory  of  respiration  can  possibly  hold 
its  ground  which  does  not  include,  nor  can  include,  the  special 
phenomena  in  the  structures  and  functions  in  the  lungs,  and 
which,  of  course,  would  include  the  nervous  arrangements  for  co- 
ordinating the  muscular  envelope  and  blood-vascular  system 
with  them,  in  order  to  produce  the  rhythmical  changes  of 
pressure  in  the  organs  for  compelling  efflux  of  air  and  blood  in 
the  alveoli  for  respiratory  purposes,  at  the  same  time  keeping 
the  fluids  in  their  channels  and  maintaining  a  balance  in  the 
circulation.  And  in  the  absence  of  a  fundamental  principle 
upon  which  to  base  the  mechanics,  it  were  utterly  impossible 
to  do  this,  as  a  matter  of  course. 

♦  But  by  applying  this  law  of  pressure  and  the  power  of  pro- 
ducing rapid  rhythmical  changes  in  pressure  to  the  special 
circumstances  in  the  structures  and  functions  in  the  lungs, 
together  with  the  principle  for  evolving  force  in  the  organism, 
and  the  whole  is  at  once  made  intelligible.  The  special 
arrangements  that  obtain  in  the  aveoli,  the  manner  the  walls 
are  hooped  in  with  elastic  tissue  fibres  to  the  framework  in  the 


PHYSIOLOGICAL    ANATOMY.  95 

lungs,  the  disposition  of  the  capillary  plexuses  upon  the  walls, 
the  residual  air  for  ballooning  them  so  as  to  obviate  friction, 
the  special  arrangements  in  the  tracheal  system  for  maintain- 
ing patency,  so  as  to  effect  free  ingress  and  egress  to  the  air, 
together  with  the  numerous  muscles  and  nerves  for  operating 
them,  the  circumstance  of  suspending  the  lungs  from  the 
upper  dorsal  vertebrae  so  as  to  hang  free  in  the  pleural  cavi- 
ties, and  entirely  disconnected  from  the  ribs,  the  purport  of 
the  pneumogastric  and  recurrent  laryngeal  nerves  ;  finally,  the 
relations  which  the  heart  sustains  to  the  lungs,  together  with 
the  special  arrangements  in  the  nerves  for  coordinating  the 
heart  and  blood-vascular  system  with  respiration  ;  all  these 
things  are  made  intelligible ;  otherwise  are  utterly  inexpli- 
cable. A  number  of  others  could  be  readily  added,  but  they 
will  come  up  as  we  go  along. 

Concerning  Adjustments  in  the  Lungs. — Commencing  with 
the  external  surface,  the  first  thing  to  note  is  the  suspension  of 
the  lungs  from  the  upper  dorsal  vertebra*,  and  clear  of  all 
connection  with  the  ribs,  so  as  to  hang  free  in  the  cavities' 
of  the  pleura*. 

Of  course,  this  fact  is  impossible  of  explanation  upon  any 
other  hypothesis  than  that  of  free  action  in  the  organs,  for 
which  this  condition  is  absolutely  essential,  enabling  them  to 
expand  outward  and  downward  during  inspiration,  and  for 
contracting  upward  and  inward,  or  toward  their  roots,  during 
expiration,  thereby  bringing  all  the  air-cells  into  active 
operation,  those  in  the  apices  with  those  in  the  bases  of  the 
organs,  since  all  perform  work  in  respiration  in  proportion  to 
the  size  of  the  capillary  plexuses.  But  in  the  absence  of  this 
power  of  expanding  and  contracting  in  the  cavities  of  the 
pleurse,  the  apices  could  not  be  brought  into  action,  the  upper 
portions  of  the  chest-cavity  being  comparatively  immovable. 
There  must  be  some  easy,  natural  method  for  doing  this  other 
than  obtains  by  reducing  pressure  at  the  sides  and  bases  of 
the  lungs,  which  would  have  the  effect  of  aspirating  all  the 
air  into  these  localities  only,  and  leaving  the  apices  perfectly 
useless,  even  admitting  that  the  old  principle  for  operating 
the  lungs  did  apply,  which  by  no  means  can  be  done.  But 
the  cells,  by  expanding  simultaneously,  would  compel  cir- 


96  PHYSIOLOGICAL    ANATOMY. 

culation  of  air  to  be  uniform  throughout  the  lungs,  otherwise 
impossible. 

Furthermore,  this  would  explain  the  existence  of  the  pleurae, 
together  with  the  copious  secretions  for  lubricating  the  organs 
so  as  to  obviate  friction  during  the  rhythmical  expansions  and 
contractions  ;  otherwise  inevitable.  Then,  again,  it  would  ex- 
plain the  rapid  resorption  of  the  pleuritic  secretions  in  order 
to  maintain  a  balance  ;  and  since  they  are  rapidly  poured  out, 
some  expeditious  and  efficacious  means  must  obtain  for  compel- 
ling resorption  to  be  in  correspondence  with  secretion  for  main- 
taining a  balance,  otherwise  impossible.  And  nothing  could 
be  more  admirable  than  the  arrangements  that  obtain  in  this 
respect  For  example,  the  lymph  channels  to  these  membranes 
open  upon  the  pulmonic  and  diaphragmatic  surfaces  by  means 
of  numerous  stomata,  while  a  dual  force  applies  for  compel- 
ling them  into  the  channels  ;  and  which  is  produced  during 
inspiration  by  the  low  pressure  in  the  lungs,  with  high  pressure 
ure  in  the  pleurae,  acting  simultaneously,  so  that  the  best 
possible  conditions  obtain  during  inspiration  for  pumping 
the  pleuritic  secretions  into  the  lymph  channels;  while  during 
expiration,  and  the  expansion  in  the  diaphragm  which  this 
produces,  throws  open  its  lymph  channels ;  at  the  same 
time  high  pressure  is  produced  in  the  pleurae  by  the  viscera  in 
the  abdomen,  which  are  compressed  into  the  chest-cavity  under 
the  action  of  the  powerful  muscles  in  the  walls,  the  chest  also 
contracting.  Thus,  during  both  inspiration  and  expiration,  a 
pulling  and  a  pushing  force  combine  for  compelling  rapid 
resorption  of  the  pleuritic  secretions  ;  and  since  they  are  con- 
cerned in  haematosis  as  well,  being  lymphoid,  the  necessity 
for  such  expeditious  method  for  producing  circulation  in  the 
sacs  will  at  once  appear  obvious. 

Of  course,  the  same  mechanics  apply  for  removing  the  morbid 
collections  resulting  from  pathological  changes  in  the  organs. 
But  the  action  in  the  lungs  may  be  impeded  by  adhesive 
inflammation,  which  is  very  common — indeed,  is  nearly  always 
the  case  in  chronic  pleurisy ;  hence  the  occasion  for  early 
surgical  interference  in  acute  pleurisy  accompanied  by  copious 
serous  effusions,  lest  the  bridles  of  lymph  which  are  likely  to 
form  between  the  surfaces  should  become  firmly  organized  and 


PHYSIOLOGICAL    ANATOMY.  97 

inhibit  the  action  in  the  lungs.  This  action  in  the  lnngs  would 
also  explain  the  friction  sonnds  in  the  early  stage  of  acnte  pleu- 
risy, from  arrest  of  the  normal  secretions  and  the  drying  of  the 
membranes  which  this  produces,  the  to-and-fro  movement  in  the 
lung  as  it  expands  and  contracts  against  the  costal  pleura  giving 
rise  to  the  friction  sounds,  or  the  same  precisely  as  obtains  in  the 
early  stage  of  pericarditis,  the  rhythmical  expansions  and 
contractions  in  the  heart  causing  the  opposite  surfaces  of  the 
pericardial  membrane  to  rub  against  each  other. 

Last,  but  not  least,  it  would  account  for  the  embarrass- 
ment to  respiration  which  is  produced  by  pleuritic  adhe- 
sions, and  why  this  should  be  in  proportion  to  the  extent 
of  the  adhesions,  and  most  dangerous  when  occurring  at 
the  bases  of  the  lungs  or  most  active  portions,  as  this  would 
inhibit  free  action  in  the  organs.  But  should  the  lungs 
be  passive,  as  alleged,  then  the  more  completely  they  were 
under  the  control  of  the  containing  walls  the  more  efficient 
they  would  be,  as  this  would  compel  response  to  every  move- 
ment in  the  latter  for  pumping  the  air  into  and  out  of  the 
organs.  On  the  contrary,  however,  it  dreadfully  interferes* 
with  respiration,  and  these  unfortunates  breathe  with  diffi- 
culty; and  when  extensive  at  the  bases  of  the  lungs,  they  perish 
speedily  by  asphyxia. 

It  follows  that  the  lungs  must  be  free  in  the  cavity  of  the 
pleurae  to  admit  of  the  rhythmical  expansions  and  contractions 
in  respiration,  arid  are  not  "passive  bags,"  to  be  "pulled 
open  and  closed  by  the  action  in  the  external  walls,"  but  that 
they  perform  an  active  role.  But  the  argument  does  not  end 
here,  only  fairly  begins,  for  overwhelming  evidence,  a  perfect 
avalanche,  is  coming.     We  now  turn  to  the  mucous  surface. 

Of  course,  the  sole  purpose  of  respiration  is  to  bring  the  air  and 
blood  into  intimate  relation  for  effecting  mutual  interchange  ; 
consequently,  the  air  must  complete  the  entire  journey  to  the 
alveoli  in  order  to  carry  in  oxygen  and  bear  out  the  waste 

*  In  a  case  of  acute  pleurisy  coming  under  my  own  observation,  occurring  in 
a  young  man,  and  suddenly  terminating  fatally,  no  other  cause  of  death  could 
be  found  save  recent  extensive  adhesions  at  the  bases  of  the  lungs ;  most  exten- 
sive in  the  right,  but  affecting  large  portions  of  the  left  pkura  also  ;  in  both. 
sides  the  diaphragm  was  adherent. 


98  LUJSTG-ACTI0N    ABSOLUTELY    NECESSARY. 

products  (inclusive  of  the  foreign  matter  carried  into  the 
organs  by  the  air),  and  which  must  be  as  rapidly  removed  for 
maintaining  the  functions  in  the  lungs.  Hence,  it  is  manifest 
that  the  ■  tidal  air"  does  not  go  half -way  to  the  lungs,  then  stop 
suddenly  against  the  "  residual  air"  as  though  this  were  an 
impediment  and  a  barrier  to  further  advance,  to  transact  its 
business  with  the  venous  blood  as  best  it  can  through  this 
obstruction  ;  at  the  same  time  to  remove  the  waste  products, 
since,  this  would  by  no  means  effect  the  objects  sought  to  be 
accomplished  by  the  mechanics,  and  less  than  this  would  come 
short  of  the  scheme  in  the  circulation  and  make  it  impossible 
to  carry  on  the  functions  in  the  lungs.  Moreover,  the  very 
mechanical  principle  which  is  involved  for  pumping  air  into  and 
out  of  the  lungs  must  inevitably  produce  a  current  through  the 
tracheal  system  and  terminal  air-cells,  since  they  expand  and 
contract  regularly  and  rhythmically,  synchronous  with  respi- 
ration. But  we  have  now  to  mention  a  number  of  facts  of  a 
most  pressing  and  urgent  nature  connected  with  the  functions 
in  the  alveoli,  which  make  it  absolutely  certain  that  there  is  such 
energetic  circulation  of  air  through  the  tracheal  system  and 
air-cells,  namely :  In  the  first  place,  carbonic  acid,  by  reason 
of  its  weight,  tends  to  accumulate  in  the  alveoli ;  and  did  not 
some  effective  means  obtain  for  promptly  expelling  it,  would 
put  an  end  to  respiration  very  speedily  by  displacing  the 
oxygen  or  preventing  its  ingress.  The  noxious  exhalations 
which  are  poured  out  as  waste  products  in  the  alveoli,  and  if 
not  as  rapidly  removed,  would  inevitably  lead  to  decomposition 
and  infection  of  the  body,  since  the  animal  matter  which  is 
contained  in  them,  together  with  moisture  and  high  tempera- 
ture in  contact  with  the  atmosphere,  must  lead  to  rapid 
decomposition  with  septic  poisoning.  Hence,  this  would  in- 
hibit "stationary  air"  in  the  alveoli.  In  short,  stationary  air 
is  stagnant  air,  and  the  functions  in  the  lungs  would  not 
admit  of  this.  And  what  a  commentary  this  upon  the  universal 
appetite  for  fresh  air — its  delightful  sensations,  its  exhilarating 
and  revivifying  effects ;  yet  never  permitting  it  to  reach  the 
alveoli  at  all.  And  were  not  the  statement  set  up  in  print,  to 
be  read  of  all  men,  one  would  scarcely  credit  it ;  but  there  it 
stands,  in  hideous  irony  of  the  beneficence  in  Nature.     But 


LUNG-ACTION    ABSOLUTELY  NECESSARY.  99 

science  can  make  progress  only  when  the  facts  are  fully  and 
completely  ascertained,  and  this  great  law  of  pressure  under- 
lying the  organism  itself  never  having  been  applied  to  the 
functions  in  the  lungs  methodically,  the  phenomena  could  riot 
possibly  be  understood  ;  hence  this  monstrous  statement,  not 
to  be  believed  henceforth  and  forever  in  this  world  of  ours. 

2.  There  must  be  arrangement  for  maintaining  cleanliness 
in  the  tracheal  system  and  air-cells  by  removal  of  the  foreign 
matter,  such  as  dust,  smoke,  etc. ,  borne  in  upon  the  tidal  air, 
and  which,  by  the  force  of  gravitation,  tend  to  the  alveolar 
floors,  and,  mixing  with  the  secretions,  would  soon  form  a 
thick  layer  of  mud  upon  the  capillaries  and  so  inhibit  the 
functions  in  the  lungs,  producing  asphyxia ;  a  danger  which 
may  be  appreciated  more  readily  after  reading  the  statement 
by  Prof.  Tyndall,*  "  that  respired  air  toward  the  end  of  ex- 
piration is  strained  of  all  dust  particles  in  the  lungs,  and  is 
absolutely  pure^  referring  to  that  coming  from  the  deeper 
portions,  or  the  last  expired.  One  may  readily  imagine, 
therefore,  the  amount  of  deposit  which  must  take  place  in  the 
lungs  from  this  source,  and  which  must  be  as  rapidly  re- 
moved. But  to  this  must  be  added  the  secretions  that  are 
poured  out  upon  the  surface  for  lubricating  it ;  together  with 
the  products  of  inflammatory  action — pneumonia,  for  example 
— in  which  the  affected  air-cells  are  literally  filled  up  with 
exudation  products  and  for  the  time  obliterated.  Hence^  to 
be  commensurate  with  this  circumstance  would  require  some 
comprehensive  and  effective  means  for  expelling  them  and 
so  relieving  the  air-cells. 

Much  stress  is  laid  upon  the  cilia  for  maintaining  cleanli- 
ness in  the  lungs;  but  independent  of  the  fact  that  these  deli- 
'  cate  hairs  are  totally  incapable  of  performing  the  amount  of 
work  which  this  involves,  the  statement  would  certainly  not 
apply  to  the  alveoli,  the  place  of  places  where  cleanliness 
should  be  maintained,  since  they  fade  out  and  disappear  alto- 
gether in  the  bronchioles.  And  in  this  connection  we  are 
forced  to  notice  a  statement  of  German  origin,  which  has  gone 
the  rounds  and  would  seem  to  be  credited,  "that  the  exuda- 
tion products  in    pneumonia    are    resorbed    and    eliminated 

*  Fragments  of  Science,  p.  161.     D.  Appleton  &  Co  ,  New  York. 


100  LUNG-ACTION   ABSOLUTELY  NECESSARY. 

through  the  special  emunctories,"  and  which  is  equivalent  to 
saying  that  the  copious  septic  materials  in  the  alveoli  are 
emptied  into  the  arterial  system  and  sent  through  the  length 
and  breadth  of  the  domain  to  infect  the  nervous  centres,  and 
form  innumerable  foci  for  inflammatory  action  ;  or,  in  other 
words,  that  the  road  to  recovery  in  pneumonia  is  by  septicaemia, 
which  is  preposterous.  Ach,  Gott !  It  is  too  horrible.  It 
will  not  do  to  think  about.  That  Nature  would  do  this 
thing!  That  she  should  sweep  the  festering  alveolar  col- 
lections into  the  blood-vessels  in  order  to  get  rid  of  them, 
when  they  are  virtually  already  out  of  the  body,  being 
upon  the  mucous  surface,  and  for  their  complete  expulsion, 
by  a  most  short,  direct  and  expeditious  route,  a  most  perfect 
mechanical  arrangement  obtains,  is,  to  say  the  least,  and 
putting  it  mildly,  a  most  extraordinary  statement.  Perceiving 
this,  therefore,  we  only  mention  the  circumstance  to  show  how 
urgent  the  necessity  for  a  further  extension  of  purely  mechan- 
ical principles  in  the  body  in  order  to  make  the  circumstances 
in  the  structures  and  functions  of  the  organs  intelligible.  In 
fine,  the  collections  are  pumped  out  of  the  alveoli  by  the 
rhythmical  expansions  and  contractions  in  the  bronchial  tub- 
ing during  respiration,  and  which  are  much  more  energetic 
than  in  the  alveoli.  The  tough  fibrinous  concretions  are  first 
detached  from  the  underlying  capillaries  and  more  or  less  dis- 
integrated by  the  purulent  secretions  which  they  provoke, 
when  they  are  readily  aspirated  by  the  bronchial  tubing  and 
expelled  thence  by  cough — a  sudden  and  forced  expiration  for 
sweeping  out  the  bronchial  collections,  upon  the  same  principle 
precisely  as  obtains  in  sneezing  for  expelling  matter  irritating 
the  Schneiderian  membrane.  It  is  easily  understood.  Thus, 
the  collections  when  aspirated  into  the  bronchial  tubing  by 
their  energetic  expansions  during  the  inspiratory  efforts,  ex- 
cites cough,  which  expels  them  ;  the  irritation  they  produce 
in  the  bronchial  mucous  membrane  being  propagated  thence 
to  the  medulla  oblongata  for  setting  up  this  reflex  action  ;  a 
circumstance  which  is  also  proven  by  exciting  the  bronchial 
mucous  membrane  artificially,  when  cough  is  readily  produced. 
And  it  would  also  explain  the  short,  energetic  inspiration 
■which  precedes  cough,  since  this  would  have  the  effect  of 


EXPELLING  ALVEOLAK  COLLECTIONS. 


101 


aspirating  the  collections  in  the  alveoli,  while  the  sndden 
forced  expiratory  effort,  which  follows  immediately  after, 
should  have  the  effect  of  expelling  them  from  the  body.  The 
anatomical  dispositions  are  very  perfect ;  could  not  be  better. 

For  example,  the  bronchi  are  surrounded  by  neighboring 
alveoli,  which  are  not  only  in  contact  with  the  tubing,  but  are 
actually  incorporated  in  the  walls  for  compelling  them  to  re- 
spond to  the  movements  in  the  tubing  during  respiration  (Fig. 
2  ,f,  <z,  b).  The  effect  of  this  arrangement  is  sufficiently  ob- 
vious. Thus,  when  the  bronchi  expand  during  inspiration, 
they  push  against  the  neighboring  alveoli  ;  at  the  same  time 
they  develop  a  suction  force  within  themselves  for  aspirating 


b     a. 


Fig.  38. — Portion  of  a  transverse  sectioD  of  a  pig's  bronchial  twig,  .04  millim.  in  diam- 
eter, magnified  2^. — Schulze.  a,  outer  fibrous  layer ;  b,  muscular  layer  ;  c,  inner 
fibrous  layer  ;  d,  epithelial  layer  ;  /,  one  of  the  neighboring  alveoli. 

the  contents  ;  hence  a  pulling  and  pushing  force  combine  for 
expelling  the  alveolar  collections  during  inspiration ;  while 
during  expiration  the  sudden  contraction  which  sets  in,  and 
the  coincident  narrowing  this  produces  in  the  lumen  of  the 
tubes,  increases  the  friction  of  the  air  against  the  membranes 
for  compelling  the  secretions  out  of  the  body,  which  is  finally 
accomplished  by  cough,  in  which  the  mucous  surface  is  swept 
as  with  a  broom  by  the  out-rushing  air,  under  the  energetic 
action  of  the  abdominal  muscles  and  the  pressure  of  the  viscera 
against  the  bases  of  the  lungs  which  this  produces,  the  sides 
of  the  chest,  of  course,  contracting  simultaneously.     And  the 


102  PHYSIOLOGICAL    REQUIREMENTS 

more  energetic  this  is  made  the  more  efficient  it  should  be  for 
expelling  the  collections.  Nothing  could  be  more  perfect,  and 
the  adaptation  of  means  to  ends  is  marvelous  in  the  extreme, 
the  crowning  point  being  its  simplicity.  But  one  thing 
calls  for  another.  Thus,  this  power  of  expanding  and 
contracting  the  bronchial  tubing,  produced  by  the  muscles 
in  the  walls  (b),  calls  for  the  deep  foldings  in  the  bronchial 
mucous  membrane  (d),  which  cannot  expand  and  contract 
with  the  muscular  walls ;  together  with  the  circular  arrange- 
ment of  the  folds  which  occurs  around  the  longitudinal  axis 
of  the  tubes,  is  in  itself  a  forceful  illustration  of  this  principle 
in  the  mechanics.  It  also  occurs  in  the  intestines,  bladder,  etc., 
but  is  by  no  means  so  great,  comparatively,  as  in  the  bronchial 
tubing  ;  in  all  which  the  containing  walls  are  composed  of  a 
basis  of  circular  muscles  and  elastic  tissue  fibres,  or  such  as  ob- 
tains in  the  worms,  and  which  has  its  explanation  in  the  fact 
of  a  common  organic  law  underlying  animal  life  for  regu- 
lating the  movements  of  the  fluids,  and  which  calls  for  similar 
arrangements  in  the  structures  for  compelling  this  circum- 
stance, as  has  already  been  remarked.  We  must  conclude 
that  these  extensive  foldings  in  the  bronchial  mucous  mem- 
brane are  indicative  of  unusual  powers  of  expansion  and 
contraction  in  the  bronchi  and  bronchioli.  Thus  we  go  along 
easily,  naturally  and  without  the  slightest  trouble  in  the 
investigations,  the  law  itself  being  simple,  while  the  anatom- 
ical adjustments  are  in  accordance  with  the  requirements. 

Finally,  nothing  is  more  common  than  catarrhal  affections 
of  the  bronchial  mucous  membrane,  in  which  great  quantities 
of  mucus  and  muco-purulent  secretions  are  poured  out  into  the 
tubes  and  air-passages,  and  did  not  some  effective  means  apply 
for  expelling  them,  speedy  asphyxia  would  be  inevitable. 

Indeed,  there  is  very  little  doubt  but  that  lobular  pneumonia 
is  produced  in  this  way,  the  secretions  being  so  abundant  and 
tenacious  in  the  affected  bronchi  as  to  resist  the  efforts  at 
expulsion,  and  by  inhibiting  the  ingress  of  air  in  the  vesicles 
leads  to  enormous  engorgement  in  the  plexuses  and  copious 
effusions  ;  but  becoming  more  and  more  purulent,  consequently 
less  and  less  tenacious,  are  finally  expelled  in  the  manner 
referred  to,  and  everything  runs  on  as  before. 


EXPELLING   FOREIGN    MATTER.  103 

To  briefly  summarize  :  Expansion  in  the  bronchial  system 
during  inspiration  is  sudden  and  energetic,  while  that  in  the 
alveoli  is  slow  and  gentle,  lasting  through  the  whole  period 
of  inspiration.  The  effect  would  be  to  aspirate  the  atmosphere 
and  the  alveolar  collections  simultaneously,  the  fluids  passing 
into  the  bronchial  system  from  both  directions ;  from  above  as 
well  as  from  below.  But  the  wave  of  expansion  passing  from 
the  bronchial  tubing  over  the  alveoli,  producing  a  slow  and 
gentle  action  in  the  latter,  causes  the  air  to  pass  gently  over 
the  collections  in  the  tubing  to  reach  the  alveoli,  not  sweeping 
them  back  again,  the  wide  expansion  in  the  lumen  causing  the 
air  to  pass  over  them  with  the  least  amount  of  friction  ;  while 
the  succeeding  contraction  during  expiration  would  have  the 
effect  of  impeding  egress  of  the  air,  and  in  proportion  increas- 
ing the  friction  against  the  mucous  surface  for  compelling  out 
the  contents. 

Thus,  nothing  could  be  more  admirable  for  maintaining 
cleanliness  in  the  alveoli  and  air  passages  than  this  beautiful 
device.  Moreover,  it  will  be  seen  from  the  nature  of  the 
mechanical  adjustments  which  obtain,  that  by  no  possibility 
can  there  be  any  quiescent  or  stagnant  air  in  the  lungs  in  the 
normal  condition  of  the  organs ;  but,  on  the  contrary,  that  it 
is  maintained  in  incessant  motion  through  the  tubing,  while 
the  alveoli  themselves  are  as  so  many  little  vortices,  where 
nothing  is  at  rest  for  a  single  moment  even,  the  old  air  rushing 
out,  carrying  with  it  the  waste  products,  and  the  fresh  air 
rushing  in,  with  its  life-giving  oxygen  ;  thus  fulfilling  the  con- 
ditions for  energetic  circulation  of  the  commerce  in  and  out  of 
the  lungs,  for  evolving  the  force  expended  in  the  organism. 
Then,  again,  the  remarkable  energy  with  which  the  blood  in 
pulmonary  hemorrhage  is  expelled  from  the  lungs  must  be 
accounted  for,  together  with  the  foreign  substances,  such  as 
water,  etc.,  that  have  accidentally  fallen  through  the  rima 
glottidis  ;  the  solid  substances  often  catching  here  from  pres- 
entation of  the  wrong  diameter  to  the  fissure,  and  so  falling 
back  again  and  again  to  the  bronchial  divisions.  There  is  tre- 
mendous effort  to  expel  them  on  the  part  of  the  tubing,  often 
repeated,  but,  unfortunately,  frequently  requiring  surgical 
procedure  from  the  increase  in  bulk  they  speedily  undergo  by 


104  FUNCTIONS   IN   THE  TRACHEA. 

reason  of  the  heat  and  moisture,  as  also  from  the  spasmodic 
action  they  produce  in  the  glottis  from  irritation. 

But  perhaps  the  most  suggestive  facts  of  the  active  role  in 
the  lungs  are  to  be  found  in  the  anatomical  dispositions,  since 
these  are  as  means  to  ends.  Of  course,  the  anatomy  must  be 
explained  and  reconciled  with  the  physiological  phenomena, 
as  cause  and  effect,  for  work  cannot  be  done  in  the  absence  of 
appropriate  arrangements,  and  the  adjustments  that  obtain  in 
the  lungs  for  special  work  are  marvelous  in  their  perfection 
and  comprehensiveness.     Beginning  with  the  trachea : 

In  the  first  place,  note  the  peculiarities  that  obtain  in  its 
several  divisions,  or  the  portions  external  to  the  lungs,  and 
those  within  the  lung-substance,  notably  the  cartilages  and 
the  muscles.  In  the  portion  external  to  the  lungs  the  car- 
tilages are  in  the  form  of  imperfect  rings  which  open  pos- 
teriorly and  are  filled  in  with  muscles,  while  within  the  lung- 
substance  they  are  thin  plates  disposed  around  the  tubing, 
but  leaving  interspaces  (Fig.  29,  2),  while  the  muscles  are 
circular  and  form  a  complete  cylinder  within  them,  and 
which  continues  to  the  very  air-cells  themselves  though  the 
cartilaginous  plates  gradually  fade  out  and  disappear  in  the 
smallest  divisions.  The  effects  of  these  arrangements  are 
obvious.  Thus,  the  strong  cartilaginous  rings  in  the  trunk 
and  primitive  divisions  of  the  trachea  function  as  a  frame- 
work of  support  to  the  organ  against  external  force  for  main- 
taining patency,  at  the  same  time  producing  elastic  move- 
ment during  inspiration,  thereby  increasing  suction  force 
correspondingly  for  aspirating  the  lungs  and  maintaining 
cleanliness.  Of  course,  the  more  energetic  tracheal  expansion 
is  made,  the  more  effective  it  should  be  in  aspirating  the 
collections,  as  must  appear  obvious.  The  cartilages  being 
thick  anteriorly,  tapering  posteriorly,  leaving  a  shoulder  for 
the  muscular  insertions  (Figs.  34  and  35),  could  not  otherwise 
than  have  this  effect  upon  the  tubing,  nor  could  these  ana- 
tomical dispositions  be  explained  by  any  other  theory  of 
tracheal  function  ;  at  any  rate,  it  explains  the  phenomena. 

The  thin  cartilaginous  plates  within  the  lungs  maintain 
patency  in  the  tubing  during  expiration,  the  firm  compression 
of  the  lungs  tending  to  destroy  this;  while  during  inspiration 


FUNCTIONS    IN   THE   TfiACHEA. 


105 


they  oppose  a  firm  surface  to  the  surrounding  alveoli  for  com- 
pelling the  collections  in  the  tubing  and  making  the  suction- 
action  more  effective  on  this  account. 


Coma 


Fig.  29. — Front  View  of  the  Human  Trachea  and  its  Smaller  Divisions. — Gray. 

The  circular  arrangement  in  the  muscles  has  plain  import, 
the  rhythmical  expansions  and  contractions  in  the  tubing  call- 
ing for  this  :  they  simply  elongate  and  contract,  with  inspira- 


106 


PHYSIOLOGICAL   ANATOMY. 


tion  and  expiration,  the  wave  of  expansion  extending  from 
above  downward  to  the  air-cells,  which  are  last  affected,  while 
the  wave  of  contraction  extends  from  below  upward.  Tn  this 
way,  an  energetic  circulation  is  produced  through  the  tubing 
for  respiratory  purposes,  and  cleanliness  is  maintained.  Of 
course,  it  also  effects  elongation  and  contraction  in  the  lungs 
during  inspiration  and  expiration,  enabling  them  to  expand 
outward  and  downward  during  inspiration,  and  to  contract 
inward  and  upward  during  expiration,  thus  producing  the  up- 


Pig.  30.— Section  through  a  Lateral  Infundibulum.—  Schulze.  From  the  lung  of  an 
adult  human  being,  after  it  had  been  filled  with  and  hardened  in  alcohol  containing 
acetic  acid,  a,  entrance  from  the  alveolar  passage  into  the  infundibulum  ;  the 
upper  margin  of  the  opening  has  been  partially  removed  by  the  section,  magnified 
^  ;  b,  b,  nuclei  of  smooth  muscular  fibres,  magnified  *£  ;  c,  c,  framework  of  elastic 
fibres. 

and-down  action  of  the  organs,  or  the  to-and-fro  movement  in 
the  long  axis  of  the  body,  for  which  suspension  by  the  roots 
from  the  spine  so  as  to  hang  free  in  the  pleural  sacs, 
together  with  the  copious  secretions  for  lubricating  them,  is 
essential ;  therefore  readily  accounted  for.  Finally,  external 
to  this  cylinder  of  muscles,  we  have  a  thick  coat  of  elastic 
and  connective  tissue  fibres  (as  obtains  in  arteries),  in  which 
the  cartilages  are  imbedded,  the  latter,  however,  fading  out 


PHYSIOLOGICAL   ANATOMY.  107 

and  disappearing  in  the  bronchioles  ;  but  not  the  elastic  coat, 
which  continues  on  to  the  terminal  air-cells,  forming  the  frame- 
work in  which  the  cups  are  placed  (Fig.  30,  c),  and  casting 
around  them  the  basket-like  fibres  for  imparting  strength  and 
firmness,  at  the  same  time  permitting  them  to  expand  and 
contract  the  same  as  the  elastic  coat  of  the  arteries,  with  which 


Pig.  31. — Mold  of  a  Terminal  Bronchus,  and  of  a  group  of  Air-cells,  moderately  dis- 
tended by  injection,  from  the  human  subject. — Robin,  a,  bronchiole  ;  6,  c,  sub- 
divisions ;  d,  other  sub-divisions,  showing  the  different  forms  of  the  terminal  groups 
of  air- vesicles  (e,  /,  g,  h,  i,  k),  and  the  relation  of  the  bronchial  tubing  to  them 
(referring  to  the  longitudinal  axis  simply),  magnified  ^.  The  rounded  and  knobbed 
appearance  of  the  groups  is  due  to  the  coalescence  of  the  borders  or  margins  of  the 
air-cells  around  the  central  axis,  which  forms  the  air-passages  and  route  of  communi- 
cation with  the  proximal  bronchial  tubing  ;  the  irregular  and  uneven  appearance  of 
which  is  due  to  distension  from  the  injection  and  imperfect  development  of  the  con- 
nective and  elastic  tissues,  and  of  the  unstriped  muscles  which  form  the  walls  of  the 
organs.  (The  preparation  was  made  from  an  infant  that  survived  its  birth  only  a 
few  days.) 

it  is  homologous.  And  nothing  could  be  more  perfect  than 
the  beautiful  adjustments  that  obtain  in  the  lungs  for  produc- 
ing the  work  connected  with  the  respiratory  processes.  So, 
then,  looking  at  the  anatomical  dispositions  in  the  bronchial 


108  PHYSIOLOGICAL   ANATOMY. 

tubing  and  air-cells,  the  circular  arrangement  in  the  muscles 
and  elastic  coats,  extending  up  to  the  very  air-cells  (Fig.  31), 
with  the  cartilages  all  open  and  arranged  around  the  longi- 
tudinal axis  throughout  the  whole  system  ;  the  nerves  for  con- 
necting them  with  respiration.  Last,  but  not  least,  the  suspen- 
sion of  the  organs  by  the  roots  from  the  spine  so  as  to  hang 
free  in  the  cavities  of  the  pleurae,  together  with  the  copious  secre- 
tions for  lubricating  them  for  obviating  friction  ;  all  regarded 
from  the  standpoint  of  the  law  of  pressure.  Can  it  be  doubted 
for  a  single  moment  that  they  perform  an  active  role  in  respira- 
tion and  vocalization  ?  Otherwise,  all  this  were  meaning- 
less. The  following  diagram  (Fig.  32)  exhibits  the  general 
form  of  the  lungs  as  they  adapt  themselves  to  the  form 
of  the  pleural  cavities,  together  with  the  relations  the  air-cells 
sustain  to  the  tracheal  system.  It  is  easy  to  perceive  how 
nervous  force  should  cause  them  to  respond  to  the  actions 
taking  place  in  the  containing  walls,  so  as  to  maintain  them  in 
close  contact  all  the  while  during  inspiration  and  expiration, 
expanding  outward  and  downward  during  inspiration,  contract- 
ing upward  and  inward  during  expiration — that  for  producing 
these  movements  the  organs  must  be  free  in  the  cavities,  and 
all  the  arrangements  that  obtain  should  be  just  as  they  are ; 
while  any  pleuritic  attachments  should  produce  corresponding 
embarrassment  to  respiration,  at  the  bases  the  worst,  for  this 
would  trammel  the  great  downward  movement  in  the  long 
axis  of  the  body  (vastly  the  more  imx^ortant),  when  extending 
up  the  sides  inhibiting  it  producing  death.  Finally,  we  would 
add  the  results  of  some  physiological  experiments  made  upon 
the  lungs,  which  confirm  this  view  of  lung-action  and  establish 
incontrovertibly  that  the  theory  of  a  negative  pressure  in 
the  pleurae  cannot  possibly  be  correct. 

Physiological  Experiments  Made  upon  the  Lungs. — Nota- 
bly, a  large,  lean  dog  (hound)  was  firmly  held  in  the  stand- 
ing position  by  several  assistants,  and,  electing  the  interspace 
between  the  sixth  and  seventh  ribs,  a  short  silver  tube,  bent  at 
right  angles  (Fig.  33),  was  passed  into  the  pleural  cavities — 
one  upon  either  side  at  the  most  external  portion  of  the  chest. 
They  were  one-fourth  inch  in  calibre  and  one  and  three-quar- 
ter inches  in  length,  with  a  flange  and  openings  for  securing 


LUNG-ACTION"    DEMONSTRATED. 


109 


them  to  the  sides  of  the  wounds  ;  while  they  were  bent  to  de- 
flect them  away  from  the  lungs,  at  the  same  time  it  made  it 
more  difficult  for  the  animal  to  extricate  them.  The  imme- 
diate effects  of  the  operation  not  a  little  surprised  me,  though 
I  had  previously  discounted  them. 

In  place  of  at  once  falling  down  in  a  state  of  asphyxia,  the 
animal  continued  in  the  standing  position,  but  taking  long 
and  deep  inspirations,  swelling  himself  out  to  a  prodigious 
size,  then  as  slowly  contracting  the  chest  again ;  doing  this 


Pig.  33. — Tracheal  System  and  Terminal  Air-Ceils,  in  diagram. — Dalton. 

from  seven  to  eight  times  per  minute,  the  whole  performance 
bearing  a  very  striking  resemblance  to  what  takes  place  after 
section  of  the  pneumogastric  nerves.  But  when  liberated,  he 
ran  round  and  round  the  room  as  though  seeking  for  means 
of  escape,  frequently  stopping  suddenly  upon  the  haunches 
to  bite  at  the  tubes,  first  at  one  side,  then  at  the  other,  evi- 
dently in  much  misery ;  then  as  suddenly  starting  up  again 
to  take  a  run  of  the  room,  but  never  going  far,  however,  with- 


110 


LUNG-ACTION    DEMONSTRATED. 


out  making  one  of  these  halts  for  relieving  himself  of  the 
instruments.  I  had  been  watching  him  thus  for  over  twenty 
minutes,  when  I  was  called  away  upon  urgent  business,  my 
friends  going  with  me,  and  when  I  returned,  which  was  fully 
three  hours  later,  being  unavoidably  detained,  I  found  the 
poor  fellow  unconscious,  lying  at  full  length  upon  his  left 
side,  and  breathing  heavily,  the  respirations  being  thirteen  per 
minute,  the  air  passing  into  and  out  of  the  right  tube  with  a 
loud,  whistling  sound;  the  left,  of  course,  was  obstructed.  First 
one,  then  the  other  tube  was  now  removed,  and  the  wounds 
hermetically  sealed  by  twisted  suture  ;  and  placing  a  basin  of 
water  near  him,  he  was  left  for  the  night,  I  had  thought  in  a 
dying  condition.  But  the  following  morning,  however,  upon 
opening  the  door,  to  my  infinite  surprise,  he  dashed  by  me 


Fig.  33. 

down  the  stairway  with  the  greatest  impetuosity,  and,  the 
hall  door  being  orjen,  he  escaped  to  the  street,  where  I  saw 
the  last  of  him  as  he  turned  the  opposite  corner.  He  cer- 
tainly lost  no  time  in  getting  away,  not  knowing  the  kindly 
feelings  and  intentions  he  had  inspired,  and  though  I  made 
diligent  search,  keeping  my  eye  upon  the  pound  and  the 
Health  Office  as  well,*  I  never  had  the  pleasure  of  seeing  him 
again,  so  that  he  scarcely  could  have  died  from  the  wounds, 
and  his  recovery  may  be  considered  as  pretty  certain.  Nor 
did  I  deem  it  necessary  to  repeat  the  operation,  since  the 
facts  established  by  it  prove  incontrovertibly  that  the  lungs 
are  active  organs  in  respiration,  and  not  mere  "passive  bags," 
to  be  "operated  by  the  chest- walls  and  diaphragm  ;"  other- 

*  This  occurred  at  Washington,  D.  C,  July  12,  1878. 


LUNG-ACTION    DEMONSTRATED.  Ill 

wise  death  by  asphyxia  should  have  been  produced  the 
instant  the  pleurae  had  been  thrown  open  for  free  ingress  of  air. 
Neither  could  the  rapid  absorption  of  the  air  for  producing 
recovery  be  explained.  It  is  obvious,  therefore,  that  this  can- 
not possibly  be  the  mode  of  respiration,  but,  on  the  contrary, 
that  the  lungs  expand  and  contract  regularly  and  rhyth- 
mically synchronous  with  the  action  in  the  chest-walls  during 
respiration. 

Furthermore,  this  would  explain  the  cases  of  recovery  from 
severe  gunshot  wounds  in  the  chest,  in  which  the  ball  had  tra- 
versed both  lungs,  coming  out  upon  the  opposite  side  and  leav- 
ing the  pleurae  open  for  the  time.  *  It  would,  moreover,  show  the 
utility  of  speedily  closing  these  wounds  for  excluding  the  air 
and  permitting  the  work  of  repair  and  restoration  to  proceed 
without  interruption.  Save  from  actual  hemorrhage,  they 
never  perish  outright  from  the  wounds,  but  from  the  nutritive 
changes  which  are  superinduced  by  them,  pleuritic  effusions, 
pneumonia,  pyaemia,  etc.,  though  respiration  may  be  greatly 
embarrassed,  especially  if  much  air  and  blood  are  present  in 
the  pleurae,  compressing  the  lungs  and  in  proportion  imped- 
ing their  action.  But  the  immediate  danger  passed  from  this 
source,  absorption  becomes  more  and  more  energetic  from  in- 
crease of  action  in  the  lungs  with  the  removal  of  the  contents 
in  the  pleurae,  as  this  admits  of  corresponding  expansion  in 
the  lungs.  The  following  experiment  will  show  how  very 
rapidly  air  is  absorbed  from  the  cavity  of  the  pleura  in  the 
normal  condition  of  the  lungs,  notably : 

A  medium-sized  dog  was  placed  under  chloroform  and  air 
forcibly  injected  into  the  right  pleural  cavity  through  the  sixth 
intercostal  space,  by  means  of  an  air-pump  and  canula,  until 
no  more  could  be  introduced  without  dangerous  violence,  when 
the  instrument  was  removed  and  the  wound  hermetically  sealed 
by  twisted  suture,  the  animal  recovering  consciousness  within 
a  few  minutes.  The  immediate  effect  of  the  operation  was  to 
greatly  hurry  respiration,  the  animal  gasping  for  breath,  the 
mouth  partially  open  and  the  tongue  protruding,  the  parts  dis- 
colored from  venous  stasis  in  the  systemic  capillaries,  and  the 
eyes  more  prominent  from  distension  of  the  intra-orbital  veins,, 

*  Cases  of  this  kind  frequently  occurred  during  our  great  Civil  War. 


112  UING-ACTION    DEMONSTRATED. 

with  a  look  of  distress  and  anxiety  in  them  painful  to  witness. 
He  was  very  restless,  going  but  a  few  feet  at  a  time,  from 
inability  to  maintain  the  erect  posture,  continually  rising  up 
and  lying  down,  his  whole  attention  seeming  to  be  concentrated 
upon  respiration,  which  was  voluntary.  Time,  3  P  m.  At  6 
p.  m.  he  was  lying  down,  but  got  up  immediately  when  I 
entered  the  room,  and  began  to  move  about,  but  evidently  was 
not  in  as  much  distress  as  at  first,  while  all  the  symptoms  had 
improved.  During  my  absence  he  had  drank  considerable 
water.  At  9  a.  m.  the  following  morning  he  was  bright  and 
active,  and  to  all  appearances  in  a  normal  condition,  apparently 
fully  recovered,  as  there  was  not  the  slightest  embarrassment 
to  respiration.  Thus,  within  the  space  of  less  than  eighteen 
hours  from  the  time  the  air  had  been  injected  into  the  pleurae,  it 
was  absorbed  by  the  lungs — i.  e.,  compelled  through  the 
alveolar  membranes  by  means  of  high  pressure  in  the  pleurae 
produced  by  the  air,  but  increased  by  the  expansile  action  in 
the  lungs.  In  other  words,  expansion  in  the  lungs  during 
inspiration  by  increasing  pressure  in  the  pleurae,  at  the  same 
time  diminishing  it  in  the  alveoli  (which  is  inevitable,  from 
the  very  nature  of  things),  acts  as  a  pushing  and.  &  pulling 
force  combined  upon  the  pleural  air,  and,  reaching  the  mucous 
surface  by  this  means,  is  finally  expelled  from  the  body  with 
the  tidal  air,  which  is  the  shortest  and  most  expeditious  route. 
Hence,  it  is  not  difficult  to  account  for  the  rapid  absorption  of 
air  in  the  pleurae,  upon  the  basis  of  the  organic  laws  con- 
trolling the  movements  of  the  fluids,  and  which  are  being  in- 
cessantly invoked  in  the  measure  of  the  physiological  require- 
ments, as  before  remarked. 

On  the  other  hand,  did  a  negative  pressure  obtain  in  the 
pleurae,  as  alleged,  all  this  were  utterly  inexplicable.  JS  ot  rapid 
absorption  in  the  pleurae  only,  but  the  entire  mechanics  for 
circulating  air  and  blood  in  the  lungs  as  well,  since  it  all  is 
based  upon  pressure  and  fluid  equilibrium,  which  is  being 
incessantly  invoked  by  rhythmical  changes  in  pressure.  It 
were  proper  to  pause  here  a  moment,  perhaps,  to  comment  upon 
this  theory  of  "a  negative  pressure  in  the  pleurae" — how  it 
came  to  be  so  widely  adopted,  considering  the  care  and  dis- 
crimination exercised  by    physiologists — before    proceeding 


LUNG -ACTION    DEMONSTRATED.  113 

further.     In  the  first  place,  some  explanation  is  better  than 
none,  and  the  bellows  principle  once  admitted,  ran  on  wheels, 
seemingly  able  to  take  care  of  itself,  the  object  being  to  get 
air  in  the  lungs  simply  ;  but  followed  to  its  logical  results,  the 
simile  is  seen  to  be  false,  since  it  should  apply  to  the  cavities 
in  the  pleurse  as  well  as  the  alveoli  ;  moreover,  it  should  apply 
to  the  blood  as  well  as  the  air,  pressure  being  transmitted 
through  the  body,  and  which  was  overlooked,  or,  at  least,  not 
critically  examined,  owing  to  the  absence  of  a  fundamental 
principle  upon  which  to  base  the  entire  mechanics  of  circula- 
tion, as  this  would  have  shown  the  necessity  for  connecting 
circulation  with  respiration,  which  would  at  once  have  laid 
bare  the  illusion ;  and  thus  having  no  sure  guide,  it  is  in- 
evitable that  error  should  have  crept  in,  not  here  only,  but 
everywhere  else  in  the  organism — error  after  error — literally 
leaving  nothing  explained.     Then,  again,  special  circumstances 
tended  to  confirm  and  rivet  the  error,  making  it  apparently  in- 
controvertible.   Notably,  collapse  of  the  lungs  when  the  chest  is 
opened,  and  as  though  they  had  been  previously  distended  by 
the  air  as  a  balloon,  but  which  is  false  in  theory,  and  false  in 
fact.     The  first,  for  the  reason  that  a  negative  pressure  in  the 
pleurse  (upon  which  the  theory  is  based)  would  inevitably 
suck  the  animal  fluids  into  the  pleurse,  as  well  as  air  into  the 
lungs  ;  hence,  cannot  possibly  be  correct,  as  has  been  already 
fully  shown. 

The  second,  for  the  reason  that  collapse  of  the  lungs  does 
not  take  place  after  rigor  mortis  when  the  chest  is  opened, 
the  lungs  maintaining  their  size  and  shape,  and  which  is  ab- 
solute proof  that  collapse  is  not  due  to  elastic  force  from  pre- 
vious distension  of  the  lungs,  but  to  a  vital  action  produced  by 
the  stimulus  in  the  air,  brought  suddenly  in  contact  with  them, 
for  elastic  force  is  unimpaired  by  rigor  mortis.  And  being 
produced  by  vital  action,  of  course  it  grows  less  and  less  till 
rigor  mortis,  which  puts  an  end  to  life  ;  otherwise  is  inexpli- 
cable. 

And  where  is  this  negative  pressure  in  birds  and  reptiles 
for  distending  their  lungs,  having  no  diaphragm  for  effecting 
it  \  And  in  the  frogs,  which  have  neither  ribs  nor  diaphragm, 
nevertheless  can  expand  their  lungs,  even  in  the  absence  of  a 


114  MOLDED   TO   THE   CHEST-CAVITY. 

tlir  oat-apparatus  for  assisting  it  f  It  will  not  do  at  all.  The 
principle  is  wrong. 

Then,  again,  consider  the  strain  to  the  lungs  which  is 
involved  in  this  theory  of  a  vacuum  in  the  pleurae  for  effect- 
ing distension — strained  open  through  all  the  expanse  in  the 
chest  for  filling  out  the  cavity,  with  yet  more  strain  added  to 
this  with  each  inspiration  and  every  degree  of  it,  to  last 
throughout  all  of  life,  from  the  first  inspiratory  act  at  birth 
to  the  last  expiratory  at  death — and  in  all  candor  it  must  be 
confessed  the  lungs  would  be  put  upon  a  "rack"  the  most 
infernal  that  human  ingenuity  could  devise  or  living  structure 
endure  ;  but  done  all  unconsciously.  It  will  not  do  to  think 
about — Ach  Gottf — not  for  a  moment  even.  Curing  pneu- 
monia by  inducing  septicemia  is  bad  enough,  but  this  is  worse. 

The  explanation  is  easy.  Thus,  during  intra-uterine  life 
the  lungs  are  molded  to  the  cavity,  the  incurvated  dia- 
phragm, as  well  as  the  sides,  taking  the  shape  of  the  excava- 
tion, broad  and  excavated  at  the  base  to  fit  the  convexity  in  the 
diaphragm,  and  tapering  up  to  the  conical  apices,  fitting  the 
cavity  out  and  out ;  and  at  birth  the  organs  expand  with  the 
chest,  the  diaphragm  at  the  same  time  contracting,  all  the 
parts  acting  simultaneously  under  nervous  force  propagated 
through  the  medulla  oblongata  by  means  of  sensory  impres- 
sions in  the  skin  ;  while  they  grow  pari  passu  with  the  walls 
and  the  increase  in  size  of  the  cavity  ;  and  expanding  and  con- 
tracting with  these,  strain  is  made  impossible.  But  when  the 
chest  is  invaded  and  air  admitted  into  the  cavity,  this  stimulus, 
acting  on  the  sensory  surface,  produces  the  energetic  and 
powerful  contraction  which  condenses  the  organs  and  draws 
them  out  of  shape  and  out  of  all  proportion  to  the  cavity 
which  they  occupy  ;  and  which  only  proves  the  contractile 
energy  in  the  organs,  but  not  that  they  are  distended  by  the 
air.  JNor  is  it  the  number  of  the  nerve3  in  the  pleural  surface 
which  produces  the  action,  but  the  fact  that  the  organs  them- 
selves are  sensory,  the  same  circumstance  being  seen  even  in 
plants,  but  is  especially  noticeable  in  sea  anemones,  which 
contract  the  fronds  energetically  the  instant  they  are  touched, 
then  expanding  them  again ;  this  in  the  entire  absence  of  nerves 
for  producing  the  actions. 


EHYTHMICAL   EXPANSION  INEVITABLE.  115 

But  you  cannot  make  the  lungs  expand  !  No  !  No  more 
can  you  make  the  leech  expand  ;  on  the  contrary,  he  con- 
tracts, and  contracts  the  more  energetically  the  greater  the 
irritation  ;  nevertheless,  he  expands  himself  readily  enough 
upon  occasion  ;  notably  when  sucking,  swelling  himself  out 
to  a  number  of  times  the  size  in  the  empty  condition  (Figs. 
15  and  17).  And  he  must  do  so  in  order  to  make  room  for 
the  blood ;  so,  likewise,  may  the  lungs  expand  of  them- 
selves during  inspiration  ;  nay,  must  do  so,  in  order  to  make 
corresponding  room  for  the  air  and  blood  which  flow  into  them, 
the  principle  being  the  same  in  both,  precisely  ;  while  the 
fluids  flow  into  the  interior  in  conformity  with  the  organic 
law  underlying  both  alike.  The  organ,  or  the  animal ;  what 
matters  it? — the  same  law  underlying  all  of  it.  And,  irre- 
spective of  the  anatomical  and  physiological  facts  referred  to, 
which  make  it  absolutely  certain  that  the  lungs  expand  and 
contract  regularly  and  rhythmically  synchronous  with  the 
action  in  the  containing  walls,  we  have  the  imperious  require- 
ments in  an  inflexible  law  compelling  them  to  do  so. 

In  conclusion:  The  curious  circumstance,  or  the  flapping 
in  and  out  of  the  relaxed  intercostal  muscles  in  the  case 
referred  to  produced  by  chloroform  narcosis  was  not  due  to 
changes  of  pressure  in  the  pleural  cavity,  but  to  changes  of 
pressure  within  the  lungs  themselves,  the  lung-tissue  pushing 
them  out  during  expiration  when  the  organs  are  tightly  com- 
pressed ;  while  they  were  pushed  in  during  inspiration  from 
the  inability  of  the  lungs  to  sustain  the  weight  in  the  atmos- 
phere which  this  invokes,  being  able  to  sustain  only  a  portion 
of  it,  and  sufficient  only  to  make  pressure  in  the  pleuree 
greater  than  in  the  alveoli,  which  the  scheme  calls  for.  It 
need  not  be  great,  but  the  lungs  must  expand  in  order  to  pro- 
duce it.  Fifteen  pounds  per  inch  is  too  much  for  the 
lungs,  but  since  several  ounces  only  is  necessary,  the  air 
flowing  in  at  the  same  time  through  the  trachea  for  assisting 
it  pari  passu  with  expansion,  the  action  is  readily  effected. 

Concerning  the  Action  in  the  Trachea.  —With  a  view  of  de- 
termining the  energy  in  the  trachea,  I  was  led  to  make  the 
following  physiological  experiments  upon  the  organ  : 

As  soon  as  a  bullock  had  been  dispatched  at  the  slaughter- 


116 


PHYSIOLOGICAL   EXPERIMENT   ON  TRACHEA. 


house,  but  before  life  had  become  extinct,  I  made  the  follow- 
ing section  of  the  trachea  (Fig.  34),  which  shows  the  condition 
before  contraction  sets  in,  which  occurs  some  minutes  later, 
from  10  to  15  only,  or  long  before  it  invades  the  other  struc- 
tures, and  which  is  also  suggestive,  since  it  would  indicate  un- 
usual energy,  in  the  organ.  It  will  be  seen  that  the  organ  is 
circular,  being  nearly  perfectly  round.     But  when  fully  COn- 


Fig.  34. — A  Transverse  Section  of  a  Bullock's  Trachea,  made  immediately  after  death, 
and  before  contraction  had  set  in.  A.  shoulder  of  the  cartilages  where  the  transverse 
muscles  have  insertion  ;  B,  extension  of  the  thin  cartilaginous  plates  to  near  the 
mesial  line  (3)  ;  1,  mucous  membrane  ;  3,  transverse  muscles  ;  3,  longitudinal  muscles. 

tracted,  it  presents  the  following  appearance  (Fig.  35),  which 
will  give  some  idea  of  the  energy  in  the  organ.*  The  con- 
traction is  enormous,  reducing  the  lumen  nearly  one-half, 
while  the  thin  terminal  plates  forming  the  posterior  portions 
of  the  cartilaginous  rings  ( A ,  B)  are  pulled  forcibly  in  contact — 
about  a  half -inch  space  separating  them  (Fig  34,  3,  B),  and  are 
bent  upon  themselves  by  the  action  in  the  muscles,  which  pull 
away  from  the  plates  and  put  the  loose  connective  tissue  (Fig. 
35,  C)  upon  the  stretch.  It  is  proper  to  remark,  that  at  the  point 
where  the  muscles  have  insertion  into  the  cartilaginous  rings 
there  is  an  offset  or  shoulder,  from  which  the  thin  cartilagi- 
nous plates  are  extended. 

*  As  these  cuts  were  copied  from  a  photograph,  they  may  be  taken  to  be 
pretty  accurate. 


PHYSIOLOGICAL   EXPERIMENT   OX  TRACHEA.  117 

Now,  then,  the  point  we  wish  to  make  is,  that  this  contrac- 
tion which  sets  in  after  death  describes  the  arc  of  movement 
during  life,  for  which  the  anatomical  dispositions  in  the  parts 
are  the  appropriate  adjustments,  and  which  is  so  obvious  that 
it  needs  no  argumentation. 

The  progressive  increase  of  the  cartilaginous  substance  in 
the  anterior  portions  of  the  rings  for  increasing  their  elastic 
force  should  make  expansion  very  energetic  for  aspirating  the 
alveoli,  while  the  degree  of  contraction  in  the  lumen  pro- 
duced by  the  muscles  would  show  the  force  in  the  organ  for 
expelling  the  air.     All  of  which  is  sufficiently  manifest. 


Fig.  35. — A  transverse  section  of  the  same,  made  a  few  minutes  after  death,  showing 
the  amount  of  contraction  in  the  cartilages  produced  by  the  action  in  the  muscles. 
A,  shoulder  of  the  cartilages  ;  B,  the  thin  cartilaginous  plates  pulled  together  and 
bent  upon  themselves  ;  C,  loose  connective  tissue  fibres  ;  1,  mucous  membrane  ;  2, 
transverse  muscles  ;  3,  longitudinal  muscles. 

But  this,  again,  brings  up  the  nervous  apparatus  for  co- 
ordinating the  structure  with  respiration  and  vocalization,  and 
which  brings  out  in  clear  relief  the  beautiful  adjustments  that 
obtain  in  this  respect,  otherwise  inexplicable.  Notably  the 
number  of  the  nerves  to  the  bronchial  tubing  sent  off  from  the 
pneumogastrics  (Fig.  36,  12,  13)  for  coordinating  the  action 
with  respiration  and  vocalization ;  together  with  the  devious 
course  of  the  recurrent  laryngeal  nerves,  which  first  descend 


118  PRINCIPLE    IN   VOCALIZATION. 

into  the  chest  to  connect  with  the  lungs  (9,  9),  ascending  thence 
upon  the  sides  of  the  trachea  to  the  vocal  cords,  for  coordi- 
nating them  with  the  action  in  the  lungs  and  trachea.  As  the 
principle  is  the  same  in  both,  we  need  only  take  up  the  action 
in  vocalization. 

Concerning  the  Action  in  Vocalization.— Oi  course,  the 
principle  in  vocalization  is  the  same  as  obtains  in  an  iEolian 
harp  ;  notably  the  sounds  are  produced  by  rapid  vibrations  in 
the  vocal  cords  from  the  rush  of  air  against  them,  only  that 
in  the  case  of  the  vocal  cords  tension  is  constantly  changing 
for  producing  the  variety  of  sound  characteristic  of  them,  and 
which  is  effected  by  means  of  nervous  force  acting  upon  the 
structures  ;  while  this  in  turn  is  determined  by  the  force 
in  the  outgoing  air,  the  two  being  always  in  correspondence 
in  the  normal  condition  of  the  organs.  This  being  true,  it 
follows  that  in  order  to  produce  the  vibrations  in  the  cords 
the  lungs  and  trachea  would  first  have  to  be  contracted 
for  forcing  out  the  air,  the  vibrations  depending  upon  this  ; 
hence,  the  fact  that  nervous  force  should  run  this  circle  in  the 
chest  before  connecting  with  the  vocal  cords ;  otherwise,  it 
would  be  putting  the  cart  before  the  horse.  In  other  words, 
the  vocal  cords  being  an  attachment  to  the  lungs,  they  must 
be  directly  connected  with  them  by  nervous  force  for  produc- 
ing correspondence  in  action,  the  one  depending  upon  the 
other. 

And  it  makes  no  difference  how  long  the  neck  may  be — e.  g:, 
the  giraffe — the  same  necessity  exists  for  this  nervous  connec- 
tion between  the  parts,  as  must  appear  obvious.  Finally,  for 
producing  the  rapid  vibrations  in  the  cords,  it  calls  for  quick, 
energetic  action  in  the  trachea  for  driving  the  air  forcibly 
through  them,  and  which  the  larger  movements  and  the  slower 
action  in  the  lungs  are  not  equal  to,  requiring  the  swifter 
movements  in  this  line  adjustment  for  producing  them  ;  more- 
over, the  force  should  be  directly  applied  upon  the  air  itself, 
for  which  the  special  anatomical  dispositions  in  the  trachea 
are  the  appropriate  adjustments,  making  it  easy  and  natural. 
So,  then,  we  can  readily  understand  why  the  recurrent  laryn- 
geal nerves  should  thus  connect  the  trachea  with  the  vocal 
cords,  since  this  is  necessary  for  producing  correspondence ; 


RECUKREjSTT  laryngeal  nerves. 


119 


and  as  the  medulla  oblongata  is  the  common  centre  of  nerv- 
ous force,  the  whole  performs  as  a  single  organ  only  for  pro- 


^V^tafeiJj  r/  w '/-fi  $!'  ■•'4.  -"■  %r 


_ti_26 


/f 


Fig.  36. — Distribution  of  the  Pneuniogastric. — Hirschfeld.  1,  trunk  of  the  left  pneu- 
mogastric ;  2,  ganghon  of  the  trunk  ;  3,  anastomosis  with  the  spinal  accessory  ;  4, 
anastomosis  with  the  sublingual ;  5,  pharyngeal  branch  (the  auricular  branch  is  not 
shown  in  the  figure  :  6,  superior  laryngeal  branch  ;  7,  external  laryngeal  nerve  ;  8, 
laryngeal  plexus  ;  9,  9,  inferior  laryngeal  branch  ;  10,  cervical  cardiac  branch  ;  11, 
thoracic  cardiac  branch  ;  12,  13,  pulmonary  branches  ;  14,  lingual  branch  of  the 
fifth  ;  15,  lower  portion  of  the  sublingual ;  16,  glossopharyngeal ;  17,  spinal  acces- 
sory ;  18,  19,  20,  spinal  nerves  ;  21,  phrenic  nerve  ;  22,  23,  spinal  nerves  ;  24,  25, 
26,  27,  28,  29,  30,  sympathetic  ganglia. 


120  ACTION   IN   VOCALIZATION. 

during  the  sounds  in  vocal  resonance,  inclusive,  of  course,  of 
the  muscles  in  the  abdomen  and  the  walls  of  the  chest. 
Furthermore,  that  the  trachea  does  perform  an  active  role  in 
vocalization  has  forcible  illustration  in  birds — notably,  insesso- 
rial  birds  (the  greatest  songsters)  possess  five  pairs  of  muscles  for 
connecting  the  bronchi  with  the  lower  larynx,  in  which  the 
vocal  cords  are  placed,  and  are  so  disposed  as  to  influence  both 
the  length  and  the  diameter  of  the  tubes  ;  while  the  number 
diminishes  with  the  power  of  song  in  other  birds,  the  Nata- 
tores  possessing  but  two,  Rasores  and  Grallatores  one  only  ; 
while  in  the  king  of  the  vultures  and  the  condor  the  vocal  mus- 
cles are  absent.  Then,  again,  in  Chelonia,  which  have  neither 
diaphragm  nor  ribs,  and  cannot  contract  the  houselike  body, 
we  have  a  special  arrangement  in  the  muscles  for  compressing 
the  lungs,  the  so-called  diaphragm  (Figs.  172,  42)  connecting 
with  this  action  in  the  trachea,  by  means  of  which  they  are 
enabled  to  produce  loud  vocal  resonance. 

But  man  possesses  the  most  extensive  provisions  for  pro- 
ducing the  energetic  action  here  referred  to,  since  the  carti- 
laginous interspaces  for  increasing  the  arc  of  movement  are 
widest  in  him  (Fig.  38,  3)  while  the  muscles  are  in  correspond- 
ence, and  undoubtedly  brought  about  as  adaptive  changes  for 
producing  the  variety  of  oral  sounds  characteristic  of  him,  as 
speech  calls  for  constant  exercise  of  the  parts.  It  will  thus  be 
seen  that  everything  is  in  correspondence  with  the  physiologi- 
cal requirements,  while  the  anatomy  falls  readily  into  line  as 
means  to  ends,  and  leaving  no  outstanding  quantity  refusing 
absorption,  furnishing  absolute  proof  in  itself  of  the  correct- 
ness of  the  premises. 


CHAPTER  VI. 

THE  MECHANICS  FOR  CIRCULATING  BLOOD  IN  THE  LUNGS,  AND 
THE  ACTION  OF  THE  HEART  AND  ARTERIAL  SYSTEM  IN  CON- 
NECTION  THEREWITH. 

Mechanics  for  Circulating  Blood  in  the  Alveoli — Anatomical  Dispositions  in  the  Walls  of 
the  Alveoli  for  Effecting  it,  or  for  Compelling  Circulation  of  Blood  in  the  Plexuses 
to  be  in  Correspondence  with  the  Circulation  of  Air  in  the  Alveoli — Extent  of  the 
Alveolar  Plexuses  and  the  Manner  They  are  Affected  by  Inspiration  and  Expiration 
— Functions  of  Residual-Air  in  this  Connection — An  Elastic  Cushion  for  Transmit- 
ting the  Force  in  the  Lungs  and  the  Muscular  Envelope  Upon  the  Plexuses  for  Com- 
pelling the  Blood  to  be  in  Correspondence  with  the  Circulation  of  Air  in  the  Alveolar 
Chambers — Manner  of  Maintaining  a  Balance  in  the  Dual  Circulations  in  the  Alveoli 
— Relations  which  the  Heart  Sustains  to  the  Pulmonic  Circulation — Mechanical  Prin- 
ciple in  the  Heart  Itself— Anatomical  Dispositions  in  the  Right  Side  Adjustments 
with  the  Functions  in  the  Lungs — Ditto,  Left  Side — Nerves  for  Effecting  Coordina- 
tion with  the  Lungs — Action  in  the  Arterial  System  Synchronous  with  Respiration 
— Nerves  for  Effecting  it — Physiological  Problem  Connected  with  the  Curves  in 
Blood-Pressure  Tracings  and  the  Curves  in  Intra-thoracic  Pressure — Traube's 
Curves — Physiological  Experiments  Proving  the  Existence  of  Rhythmical  Expan- 
sions and  Contractions  in  the  Arterial  System  Synchronous  with  the  Actions  in  the 
Heart  and  Lungs. 

In  order  to  see  through  the  marvelous  clockwork  in  respira- 
tion and  circulation,  two  facts  should  be  kept  uppermost  in  the 
mind,  not  to  be  lost  sight  of  for  a  single  moment  even,  namely : 

1.  That  respiration  is  the  means  for  evolving  force  in  the 
organism,  for  which  purpose  streams  of  air  and  blood  are 
passed  simultaneously  through  the  alveoli  by  means  of  rapid 
rhythmical  changes  in  pressure  and  a  comprehensive  tubular 
system  appropriate  to  each  communicating  with  the  alveoli  ; 
the  one  flowing  out  by  reflux  action  through  the  route  of  in- 
gress, the  other  into  the  left  side  of  the  heart  and  arterial 
system  on  its  way  to  the  cell-brood. 

2.  That  by  reason  of  inertia,  additional  force  is  needed  in 
the  blood  for  maintaining  it  in  correspondence  with  the  circu- 
lation of  air  in  the  alveoli,  or  a  measure  of  blood  for  a  measure 
of  air,  a  balance  being  maintained  between  them ;  hence  the 


122  PHYSIOLOGY   OF   ALVEOLAR  ACTION. 

f 

force-pump  in  the  heart  and  the  intimate  relations  it  sustains 
to  the  lungs,  together  with  the  innumerable  muscles  in  the 
vessels,  and  why  the  blood-vascular  system  as  a  whole  is  set 
to  respiratiou,  or  as  the  minute-hand  to  the  hour-hand,  by 
means  of  the  nerves  correlated  in  the  medulla  oblongata,  which 
functions  as  the  solar  centre  of  nervous  force  for  the  organism, 
coordinating  the  whole  mechanics  with  respiration,  to  the  end 
that  the  blood  and  air  may  be  pumped  through  the  lungs  for 
generating  force  in  the  measure  of  the  requirements  and  a  bal- 
ance be  maintained  in  the  organism ;  otherwise  impossible. 
From  that  standpoint — and  there  is  none  other — the  wholefield 
is  readily  overlooked  and  everything  made  plain  and  easily 
understood. 

The  mechanics  for  circulating  air  in  the  alveoli  have  already 
been  considered,  and  it  remains  to  show  how  the  blood  is  main- 
tained in  correspondence.  The  first  thing  in  this  connection  are 
the  anatomical  dispositions  in  the  alveoli,  as  means  to  ends, 
for  compelling  afflux  and  efflux  in  the  plexuses  with  inspira- 
tion and  expiration  simultaneously  with  the  air  in  the  cham- 
bers ;  proceeding  thence  to  the  phenomena  in  the  heart  and 
the  vessels  for  still  further  increasing  the  action,  bringing  it  up 
to  the  physiological  requirements.  The  arrangements  which 
obtain  in  the  alveoli  for  compelling  circulation  in  the  plexuses, 
with  afflux  and  efflux  of  air  in  the  chambers,  are  most  compre- 
hensive, at  the  same  time  extremely  simple,  notably  :  1.  The 
telangiectatic  plexuses  project  into  the  cavities  of  the  alveoli 
(Fig.  37),  which  at  once  secures  large  exposure  of  the  capillary 
surface  to  the  action  of  the  air,  for  effecting  rapid  interchange 
of  the  gases.  2.  The  plexuses  are  incorporated  with  the  alve- 
olar walls,  a  layer  extending  over  them;  and  which  not  only  se- 
cures them  firmly  in  position,  but  compels  them  to  respond  to  the 
action  in  the  walls,  so  that  when  the  walls  expand  for  sucking 
in  the  air  during  inspiration,  the  vessels  must  expand  simul- 
taneously, or  widen  and  elongate  at  the  same  time  for  sucking 
in  the  blood  ;  while  during  contraction  in  the  alveoli  for  forcing 
out  the  air  in  expiration,  the  capillaries  are  similarly  affected,  the 
lumen  being  contracted,  as  well  as  the  longitudinal  axis,  for  forc- 
ing out  the  blood.  Thus,  the  two  are  necessarily  in  concert,  the 
air  and  blood,  in  consequence,  rushing  into  and  out  of  the  cham- 


PHYSIOLOGY   OF    ALVEOLAE  ACTION. 


123 


bers  simultaneously ;  the  one  by  reflux  action  through  the  route 
of  ingress,  the  other  into  the  left  chambers  of  the  heart  and 
arterial  system,  as  before  remarked.  It  could  not  be  other- 
wise, in  the  very  nature  of  things.  The  plexuses  are  not 
rudely  stretched,  then— let  nothing  more  be  said  about  stretch- 
ing capillaries,  for  they  are  not  intended  to  be  stretched — but, 
on  the  contrary,  they  expand  and  elongate  with  alveolar  expan- 
sion, and  reverse  the  action  during  alveolar  contraction,  with  no 
strain  whatever  upon  them  during  these  movements.  The 
capillaries  and  walls  of  the  alveoli,  being  composed  of 
the  same  material  or  protoplasmic  substance,  are  alike 
capable  of  both  these  actions.      But    we  have  now  to  no- 


Fig.  37. — Section  of  the  Alveolar  Parenchyma  of  a  Human  Lung,  injected  through  the 
Arteria  Pulmonalis. — Schulze.  a  a,  Free  alveolar  margins  ;  b,  small  arterial  branch  ; 
c  c,  alveolar  walls  seen  in  transverse  section. 

tice  another  important  factor  in  the  mechanics  for  assist- 
ing the  capillary  circulation,  notably  the  action  in  re- 
sidual air,  which  it  would  be  difficult  to  overestimate,  since 
it  is  by  means  of  this  elastic  air-cushion  m  the  alveolar  cham- 
bers that  the  force  in  the  muscular  envelope  is  transmitted  upon 
the  plexuses  during  respiration  for  producing  the  requisite 
changes  in  pressure  upon  the  blood  itself  for  compelling  afflux 


124  THE   ROLE   OF   RESIDUAL   AIR. 

and  efflux  in  inspiration  and  expiration,  the  same  as  in  the  air 
itself,  at  the  same  time  that  it  obviates  friction  to  the  vessels  ; 
otherwise  inevitable.  Thus,  during  inspiration  the  elastic 
cushion  expands  with  the  reduction  in  pressure  which  this 
produces,  thereby  facilitating  the  expansile  action  in  the 
walls,  at  the  same  time  that  it  invites  air  and  blood  into  the 
locality  to  equalize  pressure,  each  flowing  in  through  its  re- 
spective channels  ;  while  during  expiration  the  cushion  is 
firmly  compressed  against  the  plexuses  for  compelling  their 
contents  into  the  left  side  of  the  heart,  an  equivalent  of  air  at 
the  same  time  escaping  through  the  air-passages  by  reflux  ac- 
tion, but  which  the  simultaneous  contraction  in  the  trachea 
tends  to  retard  or  regulate  so  as  to  maintain  a  balance  in  this 
respect.  In  other  words,  the  mechanical  principle  in  the 
alveoli  is  the  same  that  obtains  in  a  pump  with  an  air-cham- 
ber, only  that  in  the  present  instance  the  chamber  leaks,  as  it 
were,  letting  out  some  of  the  air  when  a  given  degree  of  press- 
ure is  reached.  But,  then,  considering  the  enormous  surface 
involved  in  the  alveolar  floors  (estimated  by  Leiberkuhn  at 
1,500  square  feet),  due  allowance  is  made  for  this  leakage,  but 
which  is  absolutely  necessary  for  maintaining  a  current  of  air 
into  and  out  of  the  compartments,  the  same  as  the  blood,  the 
object  being  to  bring  fresh  air  and  venous  blood  into  intimate  re- 
lation for  mutual  interchange,  as  before  remarked.  Finally,  by 
reason  of  the  mechanical  adjustments  that  obtain  in  the  lungs 
and  chest- walls,  a  limit  is  set  to  inspiration  and  expiration, 
in  order  that  only  a  given  amount  of  air  and  blood  can  be  taken 
in  or  given  out  at  a  time,  so  that  at  the  end  of  the  most  forced 
expiration  a  large  amount  of  residual  air  still  remains  in  the 
lungs,  which  prevents  collapse  in  the  walls  and  rude  contact 
of  opposing  surfaces ;  otherwise  inevitable.  Hence,  it  is  easy 
to  perceive  the  enormous  role  residual  air  performs  in  respira- 
tion. And  nothing  is  to  be  more  admired  than  the  compre- 
hensive arrangements  that  obtain  in  the  lungs  for  producing 
simultaneous  currents  of  air  and  blood  through  the  alveoli  and 
maintaining  correspondence  ;  but  a  common  law  underlying  it 
all  enables  this  to  be  done.  And  by  means  of  the  cardiac 
force-pump  and  the  innumerable  muscles  in  the  vascular  sys- 
tem inclusive  of  the  nervous  combinations  in  the  medulla  ob- 


PHYSIOLOGICAL   ANATOMY. 


125 


longata  for  coordinating  them  with,  respiration,  inertia  in  the 
blood  is  overcome  and  correspondence  readily  produced  be- 
tween the  blood  and  air  in  the  alveoli,  while  the  pumping 
movements  extend  from  centre  to  circumference  of  the  body 
and  from  the  lungs  to  every  tissue  territory,  the  whole  being 
intimately  connected  with  respiration  by  means  of  the  correla- 
tion of  the  nerves  in  the  respiratory  centre. 


Fig.  38. — Bronchi  and  Lungs,  Posterior  View. — Sappey.  1,1,  summit  of  the  lungs  ; 
2,  2,  base  of  the  lungs  ;  3,  trachea  ;  4,  right  bronchus  ;  5,  division  to  the  upper  lobe 
of  the  lung  ;  6,  division  to  the  lower  lobe  ;  7,  left  bronchus  ;  8,  division  to  the  upper 
lobe  ;  9,  division  to  the  lower  lobe  ;  10,  left  branch  of  the  pulmonary  artery  ;  11, 
right  branch  ;  12,  left  auricle  of  the  heart  ;  13,  left  superior  pulmonary  vein  ;  14, 
left  inferior  pulmonary  vein  ;  15,  right  superior  pulmonary  vein  ;  16,  right  inferior 
pulmonary  vein  ;  17,  inferior  vena  cava  ;  18,  left  ventricle  of  the  heart ;  19,  right 
ventricle. 

In  this  manner,  then,  a  dual  circulation  of  air  and  blood  is 
produced  through  the  lungs  for  respiratory  purposes,  the  waste 
products  flowing  out  with  the  tidal  air  for  redistribution,  while 
the  oxygenated  blood  passes  into  the  left  cardiac  chamber  and 
arterial  system  on  its  way  to  the  cell-brood,  as  before  remarked. 
The  four  wide,  short  pulmonary  veins  (Figs.  38, 13,  14,  15, 16) 
give  it  ready  egress  by  a  short  and  direct  route  to  the  auricle 


126  MECHANICAL   PRINCIPLE    IN   THE    HEART 

and  ventricle.  This  would  account  for  the  size  of  the  left 
auricular  and  ventricular  chambers,  the  amount  of  blood  pass- 
ing out  of  the  lungs  during  expiration  from  the  production  of 
high  pressure  in  the  alveoli  compelling  this  circumstance. 

Concerning  the  Action  in  the  Heart. — The  intimate  relation 
which  the  heart  sustains  to  the  lungs  has  its  explanation  in  the 
fact  that  the  organ,  like  the  lungs,  relates  to  oxygenation  for 
evolving  force  in  the  organism,  and  since  oxygen  is  imported 
through  the  blood  sent  into  the  lungs,  it  is  manifest,  for 
increasing  the  action,  that  circulation  in  the  organs  would  have 
to  be  increased  correspondingly — both  the  air  and  the  blood  ; 
but  the  latter  possessing  greater  inertia,  it  calls  for  a  force-pump 
for  effecting  correspondence  between  them  ;  hence  the  force- 
pump  in  the  heart  and  the  intimate  relations  it  sustains  to  the 
lungs,  inclusive  of  the  nerves  for  effecting  coordination  or  for 
increasing  the  diastoles  and  systoles  with  the  exigencies  in  the 
functions  in  the  lungs,  which  the  scheme  calls  for.  The  rela- 
tions it  sustains  to  the  systemic  circulation  will  come  in  pres- 
ently. But  the  first  thing  to  be  disposed  of  concerns  the 
mechanical  principle  in  the  heart  itself.  This  also  is  easily 
determined  by  the  law  of  pressure  underlying  the  organ- 
ism. Thus,  a  force-pump,  as  the  heart  is,  involves  two  ele- 
ments— a  suction  and  a  lifting  force,  acting  alternately — the 
one  involving  the  other.  The  former  is  represented  by  the 
cardiac  diastole,  for  aspirating  the  blood ;  the  latter,  by  the 
systole,  for  compelling  it  out  again,  in  order  to  produce  a  cur- 
rent ;  while  the  speed  of  the  current  thus  produced  is  depend- 
ent upon  the  energy  and  rapidity  of  the  diastoles  and  systoles, 
the  former  taking  precedence  of  the  latter. 

In  short,  the  blood  would  first  have  to  be  gotten  into  the 
chambers  before  it  could  be  forced  out  again  by  the  systole,  in 
this  manner  producing  a  stream  through  the  organ.  Further- 
more, that  diastole  produces  suction-force  is  fully  proven  by  the 
fact  that  pressure  in  the  heart  falls  below  atmospheric  pressure 
during  this  time,  while  systole,  of  course,  is  greatly  in  excess. 
For  example,  in  the  dog  the  minimum  pressure  in  the  left  ventri- 
cle varies  from  — 52  to  — 20  mm.  (mercury),  while  the  maximum 
pressure  is  about  140  mm.  In  the  right  ventricle  the  minimum 
fell  to  — 17,  while  the  maximum  rose  to  60  mm.    And  that  this 


MECHANICAL   PRINCIPLE   IN   THE   HEART.  127 

reduction  in  intra-cardiac  pressure  is  not  due  to  any  mechan- 
ical action  in  the  chest  was  demonstrated  by  opening  the  chest 
when  a  pressure  as  low  as  — 24  mm.  was  obtained  in  the  left 
ventricle.  ,<Groltz  and  Gaule,  Pfluger's  Archiv.,  xvii.  (1875), 
p.  100].  This  suction-action  in  the  heart  would  account  for 
the  danger  from  opening  the  veins  in  the  neck  during  surgical 
procedures  undertaken  in  that  locality,  without  proper  pre- 
cautions for  preventing  the  entrance  of  air  into  the  veins  under 
the  suction-action  in  the  heart,  producing  instant  death  by 
preventing  afflux  and  efflux  of  the  blood,  the  air  itself  filling 
the  heart,  which  cannot  get  rid  of  it,  and  so  causing  death. 
It  is  thus  seen  that  the  heart  performs  work  in  diastole  as 
well  as  systole;  reduction  in  pressure  compelling  this  circum- 
stance ;  while  during  systole,  when  the  action  is  reversed, 
more  work  is  performed  for  producing  lifting  force,  the  one 
necessarily  involving  the  other,  in  order  to  produce  a  stream 
through  the  organ,  as  before  remarked.  So,  then,  the  same 
principle  precisely  applies  for  the  heart  as  for  the  lungs,  the 
fluids  flowing  into  and  out  of  the  chambers  by  reason  of  rhyth- 
mical changes  in  pressure  within  itself,  produced  by  means  of 
the  rhythmical  expansions  and  contractions  taking  place  in 
its  walls  ;  only,  that  the  action  is  confined  to  the  blood  itself, 
for  reasons  already  given.  Far  be  from  us  the  wish  to  under- 
estimate the  work  performed  by  the  heart  in  the  circulation, 
but  its  importance  cannot  be  overestimated,  since  without  it 
evolution  would  undoubtedly  have  arrest  at  the  stage  in 
the  worms,  for  neither  could  correspondence  be  produced 
between  the  circulation  of  blood  and  air  in  the  alveoli,  nor 
the  high  pressure  in  the  arterial  system  for  increasing  the 
capillary  circulation.  But  we  would  not  anticipate.  We  will 
now  take  up  the  special  anatomy  in  the  organ  in  corroboration 
of  the  view  herein  set  forth. 
Concerning  the  Anatomical  Dispositions  in  the  Right  Side 

of  the  Heart  as  Adjustments  with  the  Functions  in  the 

Lungs. 

A  most  notable  circumstance  in  the  right  side  of  the  heart, 

as  adjustment  to  the  functions  in  the  lungs,  is  the  size  of  the 

ostia,  which  are  very  much  larger  than  in  the  left  side  of  the 

organ  ;    the  auriculo -ventricular  opening  especially,  but  the 


128  PHYSIOLOGICAL    ANATOMY. 

pulmonary  as  well,  which  is  considerable  larger  than  the 
aortic,  while  the  pulmonary  artery  itself  is  larger  than  the 
aorta.  * 

Another  circumstance  of  deep  import  is  the  insufficiency 
of  the  tricuspid  valves,  the  provision  for  reflux  in  the  venous 
system  as  a  safety-valve  for  the  ventricle  during  excessive  action 
in  the  lungs — e.  g.,  blowing  upon  wind-instruments,  carrying 
whole  notes,  etc. ,  when  the  blood  tends  to  accumulate  in  the  right 
side,  by  reason  of  the  high  pressure  which  prevails  in  the  alveoli 
during  this  time ;  and  were  not  some  safety-valve  thus  pro- 
vided, the  ventricle  must  evidently  be  clioked,  bringing  life  to  a 
sudden  termination  by  arresting  the  action  in  the  left  side  ; 
hence  this  circumstance.  Of  course,  some  blood  passes  from 
the  right  side  of  the  heart  through  the  lungs  during  expira- 
tion, but  is  not  sufficient  for  relieving  it,  as  is  fully  proven  by 
the  rapid  venous  suffusion  of  the  skin,  which  accompanies 
expiration  when  unduly  prolonged  from  venous  stasis  in  the 
systemic  capillaries,  the  blood  damming  back  from  the  heart  to 
the  tissue-territories.  Indeed,  partial  damming  of  the  blood  at 
the  right  side  of  the  heart  is  observable  in  ordinary  respiration 
by  watching  the  veins  in  the  neck  in  thin,  old  people  ;  but,  better 
still,  when  laid  bare  in  an  animal  by  removal  of  the  skin, 
swelling  up  and  becoming  prominent  during  expiration, 
and  then  as  suddenly  shrinking  during  inspiration,  producing 
the  so-called  pulsus  venosus.  Hence,  there  can  be  no 
doubt  whatever  of  the  fact  that  the  heart  of  itself  is 
incapable  of  maintaining  the  normal  current  of  the  blood 
through  the  lungs,  the  rapid  accumulations  in  the  right 
side  and  venous  system  during  expiration  proving  incon- 
trovertibly  that  it  falls  short  of  doing  this.  So,  then, 
this  safety-valve  in  the  right  ventricle  is  absolutely  neces- 
sary for  conserving  life.  In  death  from  asphyxia,  the  right 
chambers  are  full  almost  to  bursting,  the  enormous  accumu- 

*  According  to  Mr.  Peacok  (Pathological  Transactions,  Vol.  VI.,  p.  119),  the 
mean  circumference  of  the  right  auriculo- ventricular  opening  is  54.4  lines 
(4f£  inches),  while  the  left  is  but  44. 3  lines  (3f§  inches),  a  difference  of  nearly  one 
inch  in  favor  of  the  venous  ostium  ;  while  the  pulmonary  and  aortic  openings 
are  respectively  40  and  35.5  lines,  or  3if  and  3^  inches,  a  difference  of  nearly 
one-half  inch  in  favor  of  the  pulmonary  opening. 


PHYSIOLOGICAL    ANATOMY.  129 

lations  in  the  venous  system  preventing  the  heart  from  reliev- 
ing itself  of  the  blood,  it  is  needless  to  add  that  this  could  not 
occur  had  the  heart  the  power  to  compel  the  blood  through  the 
lungs  independent  of  the  pumping  action  in  the  latter  during 
respiration.  The  great  size  of  the  venous  ostia  facilitates 
afflux  in  the  lungs  during  inspiration,  under  the  suction-force 
which  this  produces,  while  the  heart  greatly  expedites  it,  con- 
tracting more  rapidly  during  this  time  for  increasing  its  action 
upon  the  blood,  whereby  time  is  gained  for  effecting  interchange 
of  the  gases ;  while  during  expiration  the  great  size  of 
the  pulmonary  artery,  with  the  semilunar  valves  at  the 
root,  admits  of  an  amount  of  blood  being  stored  in  the  in- 
terval, to  be  rushed  instantaneously  into  the  plexuses  the 
moment  inspiration  sets  in,  the  low  pressure  in  the  alveoli  at 
once  inviting  it ;  at  the  same  time,  pressure  in  the  artery  is 
maintained  by  the  ventricle  for  compelling  some  of  the  blood 
through  the  plexuses  to  the  left  side  of  the  heart,  which  acts 
in  concert  with  it  by  means  of  the  diastoles,  for  continuing  the 
current ;  the  blood  tending  to  accumulate  in  the  alveoli  by 
reason  of  low  pressure.  In  short,  the  size  of  the  venous  ostia 
and  pulmonary  artery,  the  reduced  size  of  the  chambers, 
together  with  the  imperfection  in  the  tricuspid  valves,  all  have 
their  explanation  as  adaptive  changes  to  the  functions  in  the 
lungs,  inclusive  of  the  increased  action  in  the  heart  during 
inspiration,  together  with  the  reduced  action  or  fall  in  frequency 
of  the  rhythms,  which  takes  place  during  expiration,  to  be 
considered  presently  in  connection  with  the  action  in  the 
arterial  system.  We  now  pass  to  the  left  side  of  the^  heart. 
Concerning  the  Anatomical  Dispositions  in  the  Left  Side  of 
the  Heart  as  Adjustments  with  the  Functions  in  the 
Lungs  and  Arterial  System. 

The  first  thing  to  note  in  the  anatomical  dispositions  in  the 
left  side  of  the  heart  is  the  size  of  the  chambers,  both  auricle 
and  ventricle  being  considerably  larger  than  in  the  right,  while 
the  walls  of  the  ventricle  are  also  thicker  (Fig.  3d,  vg.,  vd.); 
and  as  this  is  the  rule,  it  must  relate,  of  course,  to  special 
functions  in  the  organ  f«or  which  they  are  the  relative  adjust- 
ments.    The  explanation  is  easy  : 

From  the  very  nature  of  the  mechanics  in  respiration,  it 


130 


PHYSIOLOGICAL    ANATOMY. 


follows  that  during  expiration  and  the  production  of  high 
pressure  in  the  alveoli  a  large  outpour  of  blood  must 
occur  in  the  plexuses,  the  four  wide,  short  pulmonary  veins 
(Fig.  39,  13,  14,  15,  16)  giving  it  ready  egress  ;  hence  the 
great  relative  size  of  the  chambers  which  function  as  receiv- 
ing reservoirs,  But  Nature  is  utilitarian  to  the  last  degree, 
here  as  elsewhere.  Thus,  the  number  of  the  muscles  in  the 
left  ventricle,  together  with  the  augmented  capacity  of  the 
chamber,  increases  its  suction  and  lifting  force  correspond- 
ingly for  aspirating  the  plexuses  and  pumping  the  blood  into 
the  arterial  system  for  producing  the  high  pressure  in  the 


Fig.  39.— Diagram  of  the  Four  Cavities  of  the  Heart. —Bernard,  od,  right  auricle  ; 
vd,  right  ventricle  ;  og,  left  auricle  ;  vg,  left  ventricle.  The  arrows  indicate  the 
course  of  the  blood. 

latter,  the  importance  of  which  it  would  be  difficult  to  over- 
estimate. 

Then,  again,  during  inspiration,  when  blood  tends  to  ac- 
cumulate in  the  plexuses  from  low  pressure  in  the  alveoli, 
it  serves  for  continuing  the  stream,  acting  as  a  strong  suction- 
force  upon  it.  Last,  but  not  least,  during  expiration,  when 
blood  tends  to  accumulate  in  the  right  side  of  the  heart  from 
high  pressure  in  the  alveoli,  the  augmented  energy  it  gives  to 
the  outpour  acts  as  a  pulling-force  upon  the  venous  blood  for 


PHYSIOLOGICAL    ANATOMY.  131 

maintaining  a  current  through  the  lungs.  Hence,  everything 
works  together  for  the  common  good.  But,  unlike  what  ob- 
tains in  the  right  side,  the  auriculo-ventricular  opening  is  a 
closed  door  during  systole,  the  mitral  valves  fitting  closely  to- 
gether, for  the  alveolar  plexuses  must  be  protected  at  all  haz- 
ards— life's  portal,  as  it  were — and  reflux  would  pull  down 
the  very  pillars  of  the  temple  but  for  this  circumstance,  for 
everything  rests  upon  respiration.  When  reflux  does  oc- 
cur, however,  as  the  result  of  pathological  changes  in  the 
valves,  the  insufficiency  in  the  tricuspids  should  act  benefi- 
cially by  increasing  reflux  in  the  right  ventricle  ;  but  it  would 
involve  strain  to  the  alveolar  capillaries  and  inhibit  due  oxy- 
genation of  the  blood,  upon  which  the  activities  depend,  and 
leading  sooner  or  later  to  a  fatal  result  by  inducing  pulmonary 
hemorrhage  and  oedema  of  the  lungs.  It  is  needless  to  extend 
the  matter. 

Concerning  the  Nerves  to  the  Heart  and  Vessels  for  Co- 
ordinating them  with  the  Lungs. — From  what  has  preceded, 
it  follows  that  there  must  be  special  nerves  to  the  heart  and  ves- 
sels for  coordinating  them  with  the  lungs,  nerves  for  producing 
diastole  and  systole  and  regulating  the  rhythms,  so  as  to  meet 
the  special  requirements  in  lung-function,  or  the  exigencies 
in  its  circulation.  Thus,  in  the  heart  we  have  the  "inhibitor," 
so-called,  and  "accelerator"  nerves;  the  former  being  a 
dilator,  the  latter  a  contractor  nerve  for  the  heart,  while  both 
connect  with  the  respiratory  centre,  whereby  the  actions  are 
readily  coordinated  with  respiration  or  inspiration  and  expi- 
ration. The  principle  involved  in  the  mechanics  is  the  one  of 
reflex  action.  Thus,  during  inspiration  the  traction  upon  the 
nervous  filaments,  which  is  produced  by  expansion  in  the 
alveoli  and  tracheal  system  irritates  the  "contractor"  nerve 
("accelerator"),  while  the  compression  of  the  filaments  dur- 
ing expiration  excites  reflex  action  in  the  "dilator"  nerve 
("inhibitor"),  so  that  the  rapidity  of  the  rhythms  is  deter- 
mined by  the  depth  of  inspiration,  while  the  slowing  of  the 
same  (which  is  due  to  prolonging  of  the  diastoles),  by 
the  energy  of  expiration.  In  this  manner  the  action  in 
the  force-pump  is  regulated  by  the  exigencies  in  the  alveolar 
circulation  for  maintaining  a   current  through  the  plexuses 


132  DILATOR  AND   CONTRACTOR    NERVES. 

during  inspiration  and  expiration  ;  otherwise,  there  would  be 
undue  accumulation  of  the  blood  in  the  plexuses  during  inspi- 
ration from  the  low  pressure  this  produces  in  the  alveoli,  and 
undue  accumulation  in  the  right  side  of  the  heart  during  expira- 
tion, from  the  high  pressure  which  this  produces  in  the  alveoli ; 
hence  the  altered  rhythms  in  the  heart  during  these  periods, 
together  with  the  nervous  arrangements  for  effecting  them. 
That  this  is  the  principle  in  the  mechanics,  there  is  little 
room  for  doubt,  since  it  corresponds  with  the  ebb  and  flow  of 
blood  in  the  lungs  with  respiration,  which  it  tends  to  regu- 
late, so  as  to  maintain  a  current  through  the  lunge  as  nearly 
uniform  as  possible,  though  it  does  not  succeed  altogether  ; 
more  blood  flowing  out  of  the  plexuses  during  expiration  than 
inspiration.  Furthermore,  it  accords  with  the  results  of 
physiological  experiments  made  upon  the  nerves.  For  ex- 
ample, irritation  of  the  "accelerator"  nerve  increases  the 
rhythms,  by  shortening  the  diastoles,  while  irritation  of  the 
"inhibitor"  slows  them,  by  prolonging  the  diastoles,  and 
when  excessive  arrests  them  altogether,  the  heart  in  ex- 
treme diastole.  Then,  again,  in  artificial  respiration  the  in- 
jection of  air  into  the  lungs  increases  the  systole,  the  air,  of 
course,  distending  them,  and  so  pulling  upon  the  nervous 
filaments,  as  in  inspiration.  On  the  other  hand,  during 
forcible  efforts  to  expire  with  the  nose  and  mouth  occluded 
so  as  to  inhibit  escape  of  the  air,  and  thus  produce  unusual 
pressure  upon  the  nervous  filaments,  suspends  the  systole, 
by  prolonging  the  diastole,  causing  the  heart  to  intermit ;  is 
not  free  of  danger,  however,  and  should  be  practiced  cau- 
tiously. 

Finally,  we  have  the  phenomena  in  the  tracings  of  blood- 
pressure  as  further  corroborative  proof  of  this  principle  in  the 
haemal  mechanics,  since  the  elements  in  the  blood-pressure 
curve  correspond  with  the  physiological  requirements  in  the 
lungs,  as  announced  by  the  curve  of  intra-thoracic  pressure. 
The  following  diagram,  by  the  eminent  physiologist  at  Cam- 
bridge, presents  the  matter  forcibly  (Fig.  40).  It  represents 
two  tracings  made  from  a  dog,  taken  at  the  same  time.  One, 
a,  being  the  ordinary  blood-pressure  curve  from  the  carotid, 
and  the  other,  b,  representing  the  condition  of    the    intra- 


PHYSIOLOGICAL   PROBLEM   ELUCIDATED.  133 

thoracic  pressure  as  obtained  by  carefully  bringing  a  ma- 
nometer into  connection  with  the  pleural  cavity.  It  contains  a 
number  of  elements,  but  all  are  readily  explained. 

1.  We  have  the  large  undulations  in  arterial  pressure  syn- 
chronous with  respiration  (2,  2). 

2.  The  small  undulations  that  are  superposed  upon  the 
larger,  and  indent  the  surface  not  unlike  the  teeth  in  a  saw, 
only  they  are  not  so  regular,  and  which  are  produced  by  the 
actions  in  the  heart  and  arteries. 

3.  The  difference  in  the  rhythms  upon  the  opposite  sides  of 
the  large  undulations  (3,  4),  corresponding  with  inspiration 
and  expiration,  only  that  they  encroach  upon  one  another  ; 
the  rapid  rhythms  running  into  the  expiratory  period  (3,  e), 
and  the  slow  into  the  inspiratory  (4,  I,  i).  Finally,  we  have 
the  maximum  of  blood  pressure  corresponding  with  the  maxi- 
mum of  intra-thoracic  pressure  (2,  1).  Here  we  must  antici- 
pate a  little,  in  order  to  make  the  matter  fully  intelligible. 
The  grand  point  to  be  kept  uppermost  in  the  mind — not  to  be 
lost  sight  of  for  a  single  moment  even — is  to  bring  the  venous 
blood  into  the  alveoli  simultaneously  with  the  air  and  main- 
tain them  in  correspondence,  to  the  end  that  fresh  air  and 
venous  blood  may  be  kept  in  close  proximity  for  effecting 
mutual  interchange,  upon  which  everything  rests,  as  before 
remarked. 

Well,  in  order  to  do  this,  the  blood- vascular  system,  inclu-' 
sive  of  the  heart,  is  set  to  respiration  for  maintaining  cor- 
respondence between  the  circulation  of  blood  and  air  in  the 
alveoli,  by  means  of  the  vaso-motor  centre. 

Now,  then,  during  inspiration  the  arterial  system  contracts 
for  pushing  the  venous  blood  toward  the  chest-cavity,  thereby 
greatly  assisting  the  aspiratory  force  in  the  lungs  for  speeding 
the  blood  into  the  plexuses.  Hence,  the  rise  in  arterial 
pressure  during  inspiration,  together  with  the  increased 
rhythms  in  the  heart  and  vessels,  as  indicated  in  the  trac- 
ings (Fig.  40,  3);  which  is  plain  enough,  the  increased  action 
in  the  heart,  the  rise  in  arterial  pressure,  and  the  fall  in 
intra-thoracic  pressure.  But  during  expiration  and  the 
development  of  high  pressure  in  the  alveoli,  which  this  pro- 
duces, the  arterial  system  expands  for  receiving  the  blood 


134 


PHYSIOLOGICAL    PEOBLEM    ELUCIDATED. 


which  pours  into  it  during  this  time  from  the  plexuses,  thereby- 
reducing  resistance  to  the  inpour  of  blood  from  the  heart  and 
obviating  strain  to  the  organ,  since  room  must  be  made  for  the 
blood ;  hence  the  fall  in  arterial  tension,  which  continues  on 
into  the  earlier  portions  of  inspiration  (e.  i.,  2,  4,  5).  In  this 
manner,  then,  the  large  undulations  in  arterial  pressure  syn- 
chronous with  respiration  (2,  2)  are  formed,  two  of  them  pre- 
senting in  the  tracings. 


Pig.  40.—  Comparison  of  Blood-Pressure  Curve,  with  Curve  of  Intra  Thoracic  Pressure. 
To  be  read  from  left  to  right.  The  figures  are  added. — Foster,  a,  blood-pressure 
curve,  with  its  respiratory  undulations,  the  slower  beats  on  the  descent  being  very 
marked  ;  b,  curve  of  intra-thoracic  pressure  obtained  by  connecting  one  limb  of  a 
manometer  with  the  pleural  cavity.  Inspiration  begins  at  <,  expiration  at  e.  The 
intra-thoracic  pressure  rises  very  rapidly  after  the  cessation  of  the  inspiratory  effort, 
and  then  slowly  falls  as  the  air  issues  from  the  chest ;  at  the  beginning  of  the  inspira- 
tory effort  the  fall  becomes  more  rapid. 

But  while  this  action  is  going  on  in  the  arterial  system,  the 
rhythms  in  the  heart  and  arteries  are  altered  correspondingly, 
being  increased  in  frequency  during  inspiration  (3,  i,  e)  by 
hurrying  the  systoles  and  shortening  the  diastoles,  the  one 
involving  the  other,  the  object  being  to  increase  afflux  in  the 
plexuses,  so  that  time  may  be  gained  for  effecting  interchange 
before  the  succeeding  expiration  shall  have  forced  it  out  of  the 
plexuses  into  the  left  side  of  the  heart  and  arterial  system  ; 
while  during  expiration  the  systoles  are  slowed  by  prolonging 
the  diastoles  (2,  4,  e,  i),  the  object  being  to  expedite  the  flow 
into  the  left  side  of  the  heart  and  arterial  system,  at  the  same 
time  that  it  acts  as  a  pulling  force  upon  the  blood  in  the  lungs 
and  the  right  side  of  the  heart,  where  it  tends  to  accumulate 


PHYSIOLOGICAL   PROBLEM   ELUCIDATED.  135 

by  reason  of  the  high. pressure  in  the  alveoli  during  this  time  ; 
but  the  prolonged  diastoles  in  the  left  ventricle  tend  to  pull 
it  on  again  and  thus  complete  the  circuit  in  the  lungs,  the 
right  side  at  the  same  time  assisting  it  by  increasing  pressure 
upon  the  opposite  side,  while  the  whole  arterial  system  ex- 
pands in  order  to  make  room  for  the  blood  thus  surging  into 
it  during  expiration.  Hence  the  rise  and  fall  in  arterial  press- 
ure, corresponding  with  inspiration  and  expiration,  together 
with  the  rise  and  fall  in  the  cardio-arterial  rhythms  super- 
posed upon  the  large  undulations  answering  to  the  action  in 
the  heart  and  arteries.  By  increasing  diastole,  you  increase 
suction-force  correspondingly,  at  the  same  time  it  reduces  the 
rhythms.  The  discrepancies  in  the  curves  of  blood-pressure 
and  the  curves  of  intra-thoracic  pressure  are  easily  explained : 
thus  the  rise  in  arterial  pressure  during  the  early  periods 
of  expiration  (2,  1 )  is  due  to  the  sudden  damming  in  the  right 
side  of  the  heart  and  venous  system,  which  prevents  due 
escape  of  blood  through  the  capillary  system,  and  so  tending 
to  increase  pressure  in  the  arteries  ;  but  the  sudden  expansion 
which  now  sets  in  in  the  arterial  system  produces  the  sudden 
fall  in  pressure  (2,  4),  as  indicated  by  these  tracings,  and  which 
continues  on  into  the  earlier  periods  of  inspiration  (5, 6,  i),  where 
arterial  contraction  again  sets  in,  which  pushes  up  the  blood 
column  again,  as  before.  The  swell  in  intra-thoracic  pressure 
in  the  early  stage  of  expiration  (1)  is  due  to  reversal  of  inspira- 
tion by  expiration  and  the  sudden  checking  of  the  ingoing 
stream  which  this  produces,  after  which  the  stream  is  uni- 
form ;  consequently,  the  tracing  is  a  straight  line  till  inspira- 
tion is  reached,  when  another  wave  occurs  (i,  6)  indicating  the 
fall  in  intra-thoracic  pressure.  Thus,  all  the  phenomena  in 
the  tracings  are  explained  or  accounted  for,  the  whole  being 
based  upon  rhythmical  changes  in  pressure  extending  over 
the  entire  mechanics  in  respiration  and  circulation  ;  at  the 
same  time,  it  is  evident  that  Nature  nurses  the  blood  appa- 
rently with  the  greatest  care.  Nay,  more  than  this  ;  the  arte- 
ries not  only  expand  and  contract  synchronous  with  respira- 
tion, but  with  the  action  in  the  heart  as  well ;  which  we  will 
come  to  presently,  placing  the  matter  beyond  the  shadow  of 
a  doubt  or  the  possibility  of  mistake. 


136  RHYTHMIC    CENTRE   FOE   THE   VESSELS. 

Finally,  that  there  are  such  nervous  combinations  in  the 
medulla  oblongata  for  producing  the  rhythmical  contractions 
and  expansions  in  the  arterial  system  synchronous  with  res- 
piration, is  fully  proven  by  the  fact  of  their  continuance  after 
respiration  is  suspended,  "producing  the  so-called  Traube's 
curves  (Fig.  41,  2,  3),  very  similar  to  the  previous  ones,  except 
that  their  curves  are  larger  and  of  a  more  sweeping  char- 
acter." Indeed,  the  distinguished  author  fully  concedes  the 
circumstance  in  the  following  forcible  language  :*  "These  new 
undulations,  since  they  appear  in  the  absence  of  all  tho- 
racic or  pulmonary  movements,  passive  or  active,  and  are 
witnessed  even  when  both  vagi  are  cut,  must  be  of  vaso- 
motorial  origin  ;  the  rhythmic  rise  must  be  due  to  a  rhythmic 
constriction  of  the  small  arteries  due  to  a  rhythmic  dis- 
charge from  the  vaso-motor  centres,  and  especially  from 
the  medullary  vaso-motor  centre,  since  the  undulations 
are  far  less  marked  when  the  spinal  cord  is  divided.  They 
are  maintained  as  long  as  the  blood-pressure  continues  to 
rise.  With  the  increasing  venosity  of  the  blood,  however, 
both  the  vaso-motor  centre  and  the  heart  become  exhausted ; 
the  undulations  disappear  and  the  blood-pressure  rapidly 
sinks.  We  have,  then,  experimental  evidence  that,  in  the 
entire  absence  of  all  mechanical  causes,  undulations  of  blood- 
pressure,  of  direct  nervous  origin,  closely  simulating  those  ac- 
companying natural  respiration,  may  be  brought  about  when- 
ever the  blood  becomes  sufficiently  venous.  It  is  difficult  to 
imagine  why  the  vaso-motor  centre  should  exhibit  the  rhythmic 
activity  shown  in  Traube's  curves,  and  why  that  rhythm 
should  simulate  the  respiratory  rhythm,  unless  the  vaso-motor 
centre  had  been  previously  accustomed  to  a  rhythmic  activity 
synchronous  with  the  rhythmic  activity  of  the  respiratory 
centre."     Italics  are  added 

It  is  needless  to  extend  the  matter,  for  we  have  seen  that  the 
law  in  the  circulation  and  continuity  in  force  alike  call  for  this 
nervous  combination  in  the  medulla  oblongata  for  making  res- 
piration and  circulation  a  connected  movement,  in  order  to 
maintain  a  balance  between  supply  and  demand  in  the  cell- 

*  Text  Book  of  Physiology,  pp.  385,  386. 


EHYTHMIC   CENTRE  FOE  THE  VESSELS 


137 


Ibrood,  at  once  the  object  of  all  this  arrangement,  and  that  it 
could  not  be  done  in  any  other  way.  But  it  is  well  to  note, 
however,  the  physiological  adjustment  which  obtains  respect- 
ing it,  for  the  venous  blood  itself,  is  the  stimulus  for  increas- 
ing the  action  in  the  respiratory  and  vaso-motor  centres, 
whereby  venosity  in  the  blood  is  rapidly  reduced  and  the  bal- 
ance once  more  restored  when  disturbed  by  defective  respira- 


'  i  !  i 

« !  i 

Hi 


i    f\ 


t 

i  ■ 
i  ;  i 
i  \i 

i    " 


ft    r.     r-    (\    r-    !i  •"      l  i  I" 

in':  •'  !  '  !  '  ,-.'  "•/    J     / 
/  \!  \l   !  /   !/   "    "  * 


1/ 


l  i\ 
i  i  '- 
ii  ! 
if  I 
1/    '. 


I  ! 


/I   A 


ii 


WW 

i/ 

Ir     W 


\  A 

!  i  • 
!    7 


Fig.  41. — Traube's  Curves.  To  be  read  from  left  to  right. — Foster.  The  curves  1,  2,  3, 
4,  5  were  taken  at  intervals,  and  all  form  part  of  one  experiment.  Each  curve  is 
placed  in  its  proper  position  relative  to  the  base  line,  which,  to  save  space,  is  omitted. 
During  1,  artificial  respiration  was  kept  up,  the  undulations  visible  are  therefore  not 
due  to  the  mechanical  action  of  the  chest.  When  the  artificial  respiration  was  sus- 
pended these  undulations  for  a  while  disappeared,  and  the  blood-pressure  rose  steadily 
while  the  heart-beats  became  slower.  Soon,  as  shown  in  curve  2,  the  undulations 
reappeared.  A  little  later,  the  blood-pressure  was  still  rising,  the  heart-beats  still 
slower,  but  the  undulations  still  obvious  (curve  3).  Still  later  (curve  4),  the  pressure 
was  still  higher,  but  the  heart-beats  were  quicker,  and  the  undulations  flatter.  The 
pressure  then  began  to  fall  rapidly  (curve  5),  and  continued  to  fall  until  som6  time 
after  artificial  respiration  was  resumed. 

tion.  The  pumping  actions  in  the  lungs,  heart  and  vessels 
are  all  increased  correspondingly  with  venosity  in  the  blood, 
while  the  demand  for  fresh  air  for  restoring  the  balance  be- 
comes more  and  more  imperious.  In  this  manner,  then,  the 
mechanics  in  respiration  is  self-adjusting. 


CHAPTER  VII. 

AUTOMATISM   IN   THE   VESSELS,    AS   WELL    AS    IX   THE   HEART. 

Principle  in  Cardio- Arterial  Movement — The  Arterial  System  Expands  as  the  Heart 
Contracts,  and  Vice  Versa — Mode  of  Proving  this  Action  in  the  Vessels — Evidence 
in  Arterial  Tracings — Arterial  Tracings  Contrasted  with  the  Artificial  Tracings, 
Produced  by  the  Apparatus  of  Marey,  Showing  Essential  Points  of  Difference,  and 
Proving  Incontrovertibly  Automatism  in  the  Vessels — The  Variations  in  Blood- 
Pressure  Tracings  Corroborative  of  this  Action  in  the  Vessels,  Confirming  the  De- 
ductions from  Arterial  Tracings — All  the  Elements  in  Arterial  Tracings  and  the 
Tracings  which  are  Produced  by  Variations  in  Blood-Pressure  Harmonize — The 
Facts  in  Development  Confirmatory  of  Automatism  in  the  Vessels — Regular  Rhyth- 
mical Expansions  and  Contractions  Taking  Place  in  the  Vessels  of  the  Worms  in  the 
Entire  Absence  of  a  Heart  for  Producing  Them,  the  Vessels  Expanding  and  Con- 
tracting Themselves — The  Pulsations  in  the  Umbilical  Cord  not  Synchronous  with 
the  Action  in  the  Foetal  or  Maternal  Heart  ;  Moreover,  Pulsates  even  after  Connec- 
tion is  Severed — The  Pulsations  in  the  Ears  of  a  Rabbit  and  in  the  Wings  of  a  Bat 
not  Synchronous  with  the  Action  in  the  Heart,  Showing  Independent  Action  in  the 
Vessels — The  Special  Anatomy  in  the  Vessels  Fundamentally  the  Same  as  in  the 
Heart — Progressive  Increase  of  the  Muscles  in  the  Arterial  System  from  the  Heart 
to  the  Tissue-Territories — Function  of  the  Strong  Elastic  Coat — How  High  Pressure 
is  Produced,  and  the  Occasion  for  it — How  Reflux  in  the  Capillaries  is  Obviated — 
Mode  of  Increasing  and  Diminishing  the  Local  Actions,  and  for  Coordinating  the 
Vessels  with  Respiration  and  the  Action  in  the  Heart. 

The  arteries  perform  a  much  higher  role  in  the  mechanics 
of  circulation  than  as  conduits  simply,  expanding  and  con- 
tracting upon  their  contents  for  increasing  circulation,  the 
same  as  the  heart,  the  principle  being  the  same  Further- 
more, the  actions  alternate,  the  vessels  expanding  as  the 
ventricle  is  contracting,  and  vice  versa,  the  same  applying 
for  both  sides  of  the  heart.  The  following  are  the  reasons  upon 
which  this  statement  is  based  :  (a)  By  the  arteries  expand- 
ing synchronously  with  contraction  in  the  ventricles,  it  would 
have  the  effect  of  reducing  resistance  to  the  m-pour  of  blood 
from  the  ventricles,  and  in  proportion  obviating  strain  to  the 
heart  and  rude  force  to  the  blood  itself,  the  blood  corpuscles 
especially  being  too  delicate  to  withstand  it.  (b)  It  would 
explain  numerous  phenomena  anatomical  and  physiological, 


AUTOMATISM  IN"   THE  VESSELS.  139 

appertaining  to  the  vessels  ;  otherwise  inexplicable,  (c)  Last, 
bnt  not  least,  this  automatism  in  the  vessels  corresponds  with 
the  facts  in  development,  notably  the  existence  of  inde- 
pendent pulsations  in  the  vessels  in  the  entire  absence  of  a 
heart  for  ^producing  them,  the  vessels  expanding  and  con- 
tracting themselves. 

For  the  sake  of  simplicity,  we  begin  with  the  arterial  trac- 
ings as  indicative  of  automatism  in  the  vessels,  and  the 
manner  they  are  coordinated  with  the  heart  and  with  respira- 
tion ;  proceeding  thence  to  the  special  anatomy  in  the  organs 
as  means  to  ends,  or  the  provisions  for  special  work  per- 
formed by  the  vessels ;  and  concluding  with  the  facts  in 
development,  for  all  these  circumstances  should  be  reconciled 
and  explained,  thereby  proving  the  correctness  of  the  premises. 
If  a  given  theory  of  the  animal  circulation  cannot  account  for 
these  circumstances,  it  is  proof  in  itself  that  it  is  false  and  un- 
reliable, not  to  be  depended  upon  for  a  single  moment  even. 

Conceiving  the  Phenomena  in  Arterial  Tracings. — There 
are  three  elements  in  arterial  tracings  :  or,  diastole,  systole  and 
the  wave  of  dicrotism.  The  first  corresponds  with  the  straight 
perpendicular  line  (Fig.  42,  b,  c),  which  occurs  during  systole 
in  the  left  ventricle,  the  vessel  expanding  against  the  button 
of  the  sphygmographic  lever,  and  so  producing  this  straight 
perpendicular  line  in  the  tracing.  The  second,  or  oblique 
crooked  line,  c,  d,  which  joins  the  first  at  an  acute  angle,  cor- 
responds with  systole  in  the  vessels,  in  which  the  button  of  the 
sphygmographic  lever  falls,  producing  this  tracing  ;  while  the 
croolc  itself  is  the  wave  of  dicrotism  passing  along  the  vessels 
during  systole. 

Thus,  there  are  three  elements  in  the  tracings  of  arterial 
action.  According  to  received  opinion,  however,  this  action  in 
the  arteries  is  believed  to  be  produced  by  the  action  in  the 
heart,  but  it  cannot  be  sustained,  the  facts  in  the  case  inhibit- 
ing it.  Indeed,  the  very  efforts  that  are  made  to  sustain  this 
view  end  by  furnishing  the  most  indisputable  evidence  of  its 
utter  fallacy. 

For  example,  Professor  Marey,  under  the  impression  "that 
the  heart  is  the  force  in  the  circulation,  and  produces  the 
action  in  the  pulse,"  had  constructed  an  apparatus  for  repro- 


140  AUTOMATISM   IN   THE   VESSELS. 

during  this  action  artificially  in  elastic  tubing,  but  which  is  no 
more  like  the  action  taking  place  in  the  arteries  than  day  is  to 
night.  And  how  such  an  impression  could  have  been  produced 
or  have  gotten  abroad  is  explicable  only  upon  the  circumstance 
of  prejudice,  or  the  mental  bias  produced  by  the  teachings  of 
tradition,  and  which  seizes  upon  every  fact  that  apparently 
sustains  it,  shadowy  and  unreal  however  that  may  be,  and  this, 
too,  in  the  acutest  intellect ;  for  stronger  evidence  of  automatism 
in  the  vessels  than  that  very  experiment  affords,  it  would  be 
difficult  to  find,  as  I  shall  now  proceed  to  show.  Fortunately, 
however,  much  evidence  of  a  totally  different  kind,  not  open 
to  controversy,  being  visible  to  the  unassisted  eye,  is  ready  to 
our  hand. 

But  concerning  this  experiment.  Briefly  summarized,  it 
consists  as  follows  :  The  bulb  of  a  gum  elastic  syringe  (Pig.  43, 
B)  is  taken  to  represent  the  left  cardiac  ventricle,  the  same,  of 


Fig.  42. — Arterial  Tracings,    b,  c,  answers  to  diastole  ;  c,  d,  to  systole,  in  the  vessels. 

course,  applying  for  the  right,  with  valves  for  obviating  reflux : 
the  tubing  upon  the  left,  with  the  end  in  the  basin  of  water,  the 
pulmonary  veins  and  the  blood  in  the  alveolar  plexuses,  that 
upon  the  right— arranged  in  three  coils  upon  an  upright  piece 
holding  as  many  sphygmographs  (Z,  Z,  Z),  to  which  they  are 
adjusted  for  registering  the  actions  taking  place  in  them,  and 
which  are  duly  recorded  upon  a  paper  of  a  revolving  cylinder 
(O) — the  arterial  system,  which  is  supposed  to  be  dis- 
charging into  the  capillary  system,  in  which  the  vessels  termi- 
nate ( V).  Thus  constituted,  the  bulb  is  now  rhythmically 
compressed  by  the  hand  in  imitation  of  the  action  in  the  left 
cardiac  ventricle.  Pardon  us  !  But,  indeed,  we  are  brought  to  a 
standstill,  unless  you  freely  allow  considerable  suction-action  in 
the  heart  for  compelling  the  blood  into  itself  in  imitation  of  the 
bulb  and  the  water,  else  the  comparison  is  false.  Pumps  may 
differ,  but  the  principle  never  changes,  and  this  it  would  be  well 
to  bear  in  mind— not  to  be  lost  sight  of  for  a  single  moment,  else 


AUTOMATISM    IN    THE  VESSELS   DEMONSTRATED. 


141 


all  is  chaos  in  this  mechanics.  Well,  the  result  of  the  three 
tracings  thus  produced  is  to  be  seen  in  the  three  undulating 
lines  (Fig.  44,  1,  2,  3).  In  the  lowest  tracing — the  one  near- 
est the  bulb  of  the  syringe  ( I ) — the  undulations  are  greatest, 
while  in  the  uppermost  tracing  (s)  they  are  smallest  and  upon 


Pig.  43. — Marey. 

the  point  of  extinction.  N"ow,  then,  where  is  the  correspond- 
ence spoken  of  as  subsisting  between  the  tracings  produced 
by  this  artificial  scheme  and  the  tracings  which  are  produced 
by  the  action  in  the  arteries  %  For  the  life  of  me,  I  cannot 
perceive  where  it  comes  in.  No,  not  at  any  place  whatever 
can  it  be  seen  in  any  one  of  these  tracings.     For  example, 


Fig.  44. — Marey. 

take  that  first  ascending  line  in  the  tracing  nearest  the  bulb, 
where  the  undulations  are  largest,  and  where  impact  of  the 
liquid  against  the  walls  of  the  tubing  is  greatest,  and  com- 
pare it  with  this  first  perpendicular  line  in  the  arterial  tracings,, 
which  corresponds  with  arterial  diastole  (Fig.  42,  b,  c):     The 


142  AUTOMATISM    IN    THE   VESSELS   DEMONSTRATED. 

former  is  not  perpendicular  at  all,  the  nearest  approach  approxi- 
mating 70  or  75  degrees  only,  while  it  rapidly  tumbles  down,  in 
i  he  third  tracing  being  upon  the  point  of  extinction  ;  whereas, 
in  the  case  of  the  arterial  tracings  this  line  is  perpendicular, 
and  remains  to  the  end  perpendicular,  even  to  the  very  capil- 
laries themselves  (Fig.  51,  V).  Thus,  two  objections  which  are 
fatal,  to  this  argument  are  at  once  made  apparent.  The  reason 
the  large  undulations  in  the  first  tracing  so  rapidly  tumble 
down  in  the  low,  flat  forms  in  the  third  tracing  is  easily  un- 
derstood, notably  :  With  every  impact  of  the  waves  against  the 
tubing  they  part  with  some  of  the  initial  force,  and  in  conse- 
quence they  rapidly  become  less  and  less  until  the  point  of 
extinction  is  reached  They  must  do  so  in  the  very  nature  of 
things.  But  in  the  arterial  tracings  the  matter  is  quite  differ- 
ent. Here  the  line  answering  to  the  initial  force  is  perpen- 
dicular at  tlie  start,  and  there  is  no  falling  away  from  this 
anywhere,  but  it  is  fully  maintained  throughout  the  limits  in  the 
arterial  system.  Nor  does  it  differ  in  the  smallest  arterioles 
from  that  which  occurs  in  the  aorta  itself.  And  how  other- 
wise account  for  this  circumstance,  but  that  the  force  in  the 
wave  sent  out  by  the  ventricle  is  supplemented  by  the  force 
inherent  in  the  walls  of  the  arteries  themselves  ?  In  other 
words,  that  the  vessels,  in  place  of  withdrawing  force  from  the 
waves,  and  thereby  crushing  them,  as  in  example  of  the  arti- 
ficial scheme,  expand  of  their  own  accord  in  concert  with  the 
waxes  from  the  heart ;  and  by  thus  producing  a  vis  a  fronte, 
the  blood  flows  with  augmented  speed  through  the  vessels, 
with  the  least  possible  friction  upon  it,  since  resistance  is 
reduced  to  a  minimum.  Indeed  a  suction  force  is  added. 
And  this  force,  being  inherent  in  the  vessels,  should  be  co- 
extensive with  the  arterial  system  ;  hence  the  maintenance 
of  the  type-tracings  throughout  the  limits  in  the  arterial 
system ;  together  with  the  energy  in  the  arterial  pulsations, 
which  is  the  product  of  this  dual  force  in  the  vessels,  produced 
by  the  heart  and  the  vessels  themselves.  Their  strike  is 
like  a  hammer  against  the  finger-tips,  while  the  perpendicular 
line  which  this  produces  in  the  tracings  will  give  some  idea  of 
the  energy  in  this  expansile  action. 
In  reference  to  that  second  line  (Fig.   42,   c,   d),  in  the 


AUTOMATISM    IN   THE   VESSELS   DEMONSTBATED.  143 

arterial  rhythm  which  answers  to  systole  in  the  vessels, 
and  in  which  the  dicrotic  wave  presents  :  Where  is  resem- 
blance to  be  found  between  this  and  the  corresponding  portion 
of  the  tracings  in  the  artificial  scheme?  Not  to  be  found, 
either  !  No.  not  the  faintest  trace  in  anyoi  the  tracings  of  the 
dicrotic  wave  which  is  superposed  upon  it,  producing  the 
small  wave  or  undulation  in  this  line.  Nor  can  it  be  produced 
artificially,  since  it  involves  independent  or  automatic  action 
in  the  vessels  themselves.  And  in  the  absence  of  the  cardinal 
circumstances  in  arterial  movement,  one  may  not  speak  of 
correspondence  at  all.  Hence  we  must  conclude  that  that 
artificial  scheme  which  is  based  upon  vis  a  tergo  simply,  does 
not  represent  the  principle  in  car  do -arterial  action  which 
combines  vis  a  fronte  with  vis  a  tergo,  acting  simultaneously 
and  which  accords  with  the  special  phenomena  in  the  trac- 
ings, the  phenomena  in  the  heart  and  arteries,  anatomical  and 
physiological,  and  satisfies  the  physiological  requirements  in 
the  blood  itself,  and  the  blood-supply  to  the  organs.  What 
amount  of  force  Professor  Marey  applied  to  the  bulb  in  imi- 
tation of  the  cardiac  systole,  we  are  not  prepared  to  say,  but 
if  he  used  the  force  in  his  hand  he  certainly  put  as  much 
force  upon  this  artificial  circulation  as  exists  in  the  left 
cardiac  ventricle,  since  in  actual  weight  and  number  of  the 
muscles,  the  flexors  in  the  forearm  and  hand  far  exceed  the 
muscles  in  the  left  ventricle. 

But  notwithstanding  this,  the  perpendicular  line  is  not  pro- 
duced, while  the  high  undulations  in  the  early  portions  of  the 
tubing  rapidly  tumble  down  into  the  low,  flat  forms  preceding 
final  extinction  ;  as  already  remarked,  cannot  produce  either 
of  the  lines.     That  ends  the  matter. 

We  now  proceed  to  other  evidence  of  automatism  in  the 
vessels,  founded  upon  sphygmographic  tracings  also. 

The  Variations  in  Blood-Pressure  Indicative  of  Autom- 
atism in  the  Vessels. — In  the  case  of  the  arterial  tracings, 
the  actions  in  the  vessels  are  directly  reported  through  the 
sphygmograj)h  ;  whereas,  in  the  case  of  the  variations  in 
blood-pressure,  the  actions  are  communicated  to  the  sphygmo- 
graph  through  the  oscillations  taking  place  in  the  blood  itself, 
and  communicated  through  a  hsemadynamometer  to  the  lever 


144  AUTOMATISM    IN    THE  VESSELS   DEMONSTRATED. 

of  the  sphygmograph,  thereby  furnishing,  as  it  were,  the 
reverse  side  of  the  matter. 

Let  us  see,  then,  how  it  is  here,  for  this  is  important  evi- 
dence. 

We  begin  with  permanent  high  pressure  in  the  arteries  or 
zero,  which  answers  to  the  horizontal  line  in  the  tracings  (Fig. 
45),  while  the  waves  {a,  c)  represent  the  oscillations  which 
occur  in  it ;  the  large  waves  (&,  a)  corresponding  with  the 
action  in  the  heart,  the  small  ones  (c,  c)  answering  to  dicro- 
tism.  The  great  rise  in  pressure,  as  indicated  by  the  per- 
pendicular liue  upon  the  left  side  of  the  large  waves,  is 
produced  by  the  systole  in  the  vessels  ;  while  the  fali  in  press- 
ure, indicated  by  the  perpendicular  line  upon  the  right  side 
of  the  waves  (a,  b),  is  the  result  of  expansion  or  diastole,  the 
blood  column,  in  consequence,  falling  down  in  the  tube  to 
which  the  lever  of  the  sphygmograph  is  adjusted,  and  so 
registering  the  action.  But  the  circumstance  to  which  special 
attention  is  directed  is,  that  this  sudden  fall  in  pressure  ex- 
ceeds or  goes  beyond  zero,  coining  considerably  below  the  hori- 
zontal line  (Fig.  4.~,  b).  And  since  this  occurs  synchronously 
with  the  systole  in  the  heart,  is  it  not  absolute  proof  of  expan- 
sion in  the  arterial  system  during  this  time  \  Of  course,  the 
sudden  fall  in  pressure  must  be  due  to  expansion  in  the 
vessels  and  the  loss  of  support  to  the  blood-column  which 
this  occasions,  causing  it  to  fall  in  the  instrument  correspond- 
ingly ;  since  it  could  not  be  accounted  for  in  any  other  way. 
Hence,  the  more  the  arterial  expansion,  the  greater  will  be  the 
fall  in  the  blood  column,  as  must  appear  obvious.  During 
this  fall  in  pressure,  however,  the  heart  empties  itself  into  the 
arterial  system;  hence  strain  is  obviated,  and  friction  to  the 
blood  is  reduced  to  a  minimum,  the  importance  of  which  it 
would  be  difficult  to  overestimate. 

In  fine,  the  cardiac  systole  disappears,  and  is  swallowed  up 
by  the  greater  movement  taking  place  in  the  arterial  system 
itself,  in  which  it  is  merged,  as  indicated  by  the  perpendicular 
line  in  the  arterial  tracings  corresponding  with  expansion  in 
the  vessels  (Fig.  42,  5,  c). 

There  are,  then,  rhythmical  expansions  and  contractions  in 
the  arterial  system  synchronous  with  the  action  in  the  heart. 


AUTOMATISM    IN    THE  VESSELS   DEMONSTRATED.  145 

the  one  alternating  with  the  other.  In  other  words,  there  is 
complete  rhythm  in  both — in  the  arteries  as  well  as  in  the 
heart.  The  fact  must  be  borne  in  mind,  also,  that  only  one-half 
of  the  cardiac  rhythm,  or  the  systole  simply,  can  produce  any 
effect  in  the  arteries,  since  the  ventricular  diastole  can  accom- 
plish nothing  with  the  semi-lunar  valves  closed,  for  obviating 
reflux  in  the  ventricle,  as  is  the  case  in  diastole.  Now,  then. 
by  the  monistic  theory  of  the  circulation,  or  a  vis  a  tergo 
simply,  how  is  it  possible  to  explain  the  phenomena  in  these 
tracings  ?     The  thing  is  utterly  impossible. 

A  moment' s  reflection  will  make  this  obvious.  For  example : 
How  can  the  cardiac  systole  produce  the  sudden  fall  in  press- 
ure below  zero  (Fig.  45,  a,  b)  %  If  the  rise  in  pressure  {a,  a)  is 
to  be  explained  by  the  inrush  of  blood  from  the  ventricle,  how, 
then,  is  this  opposite  movement,  or  fall  in  pressure,  to  be 


Fig.  45.— The  Variations  in  Blood-Pressure  in  the  Carotid  of  a  Horse.— Marey.  The 
horizontal  line  between  the  undulations  represents  the  permanent  high  pressure  in 
the  arterial  system,  or  zero  of  pressure  in  the  arteries,  while  the  undulations  repre- 
sent the  variations,  a,  a,  the  rise  in  pressure  produced  by  systole  in  the  vessels,  and 
which  corresponds  with  diastole  in  the  heart ;  a,  b,  the  fall  in  pressure  which  answers 
to  diastole  in  the  vessels,  and  corresponds  with  systole  in  the  heart ;  c,  the  dicrotic 
wave. 

accounted  for  1  Surely  not  by  "resiliency  in  the  ve-sels/'  so 
called,  for  this  should  produce  still  higher  pressure  from  the 
contraction  it  would  occasion  in  the  vessels,  though  the  term 
itself  is  inapplicable  to  vital  action.  At  any  rate,  it  should 
have  directly  the  opposite  effect  to  that  ascribed — increasing, 
in  place  of  decreasing,  blood-pressure. 

Then,  again,  how  is  it  possible  to  produce  the  fall  in  pressure 
by  closing  the  semi-lunar  valves  simply,  as  occurs  at  the  end 
of  systole  ?  Since  the  blood  is  still  in  the  arteries ;  and  allow- 
ing for  an  amount  passing  out  at  the  distal  end  in  the  capilla- 
ries, the  contraction  or  "  resiliency ' '  would  more  than  make  up 
for  the  lo^s  in  pressure  this  tends  to  produce,  reflux  being 


146  AUTOMATISM    IN    THE  VESSELS   DEMONSTRATED. 

obviated  by  the  closing  of  the  valves.  At  any  rate,  the  fall  in 
pressure  should  be  gradual,  whereas  it  is  sudden,  abrupt  and 
extreme,  sending  it  below  zero  (Fig.  45  a,  b),  the  line  of  per- 
manent pressure ;  hence,  there  must  be  arterial  expansion  in 
order  to  effect  it,  and  that  would  very  readily  account  for  it ; 
otherwise  is  inexplicable. 

'I  his  circumstance,  then,  is  sufficient  to  show  that  the 
action  in  the  heart  would  not  of  itself  account  for  the 
phenomena  in  the  blood-pressure  tracings ;  and  taken  in 
connection  with  the  facts  already  developed  in  the  arterial 
tracings,  namely,  that  it  cannot  produce  the  perpendicular 
line  answering  to  arterial  diastole  (Fig.  42,  b,  c),  nor  the 
crooked  oblique  line  (c,  d)  answering  to  systole,  amounts 
to  actual  demonstration  of  automatism  in  the  vessels  them- 
selves, by  means  of  which  only  could  the  phenomena  be  pro- 
duced, inclusive,  of  course,  of  the  action  in  the  heart,  with  which 
the  vessels  are  coordinated.  One  other  circumstance,  how- 
ever, remains  for  mention  in  this  connection,  showing  the  cor- 
respondence subsisting  between  the  tracings  which  are  pro- 
duced by  the  oscillations  in  blood-pressure  and  arterial 
tracings,  notably  the  wave  of  dicrotism.  For  example, 
dicrotism  occurs  in  the  oblique  line  answering  to  systole  in  the 
vessels  (Fig.  42,  c,  d) ;  in  other  words,  is  produced  during 
systole,  and  which  corresponds  with  the  wave  of  dicrotism, 
which  is  produced  during  the  rise  in  blood-pressure  (Fig.  45, 
Z>,  c),  which  is  also  the  result  of  systole  in  the  vessels,  show- 
ing conclusively  that  they  are  expressions  of  the  same  thing. 
Thus,  all  the  characteristic  elements  in  the  arterial  tracings 
are  reproduced  in  the  tracings  of  blood-pressure,  the  oscilla- 
tions in  wThich  correspond  with  the  rhythmical  expansions  and 
contractions  taking  place  in  the  vessels  which  alternate  with 
the  actions  in  the  heart,  inclusive,  of  course,  of  the  movements 
synchronous  with  respiration.  In  fine,  the  mean  pressure  in 
the  vessels  is  produced  by  the  action  in  the  heart,  but  the 
oscillations  in  pressure  are  mainly  produced  by  the  vessels 
themselves,  which  have  adjustment  with  the  heart  and  with 
the  lungs,  expanding  and  contracting  upon  their  contents  to 
meet  the  special  emergencies  in  the  functions  in  these  organs  ; 
while  the  rich  nervous  plexuses  which  emboss  the  vessels  con- 


AUTOMATISM    IN    THE  VESSELS   DEMONSTRATED.  147 

necting  them  with  the  cardiac  and  respiratory  centres  serve 
for  coordinating  and  unifying  the  actions  throughout,  at  the 
same  time  also  serving  to  produce  the  local  actions  in  response 
to  special  stimulus.  Hence  the  existence  of  this  enormous 
web  of  nerves. 

But  to  this  must  be  added  the  evidence  which  is  furnished 
in  the  tracings  produced  by  changes  in  volume  (Fig.  5 1 ),  in 
which  entire  organs,  inclusive  of  the  whole  soft  tissues  in  the 
body,  expand  and  contract  regularly  and  rhythmically  syn- 
chronous with  the  actions  in  the  heart  and  in  the  lungs. 
And  to  say  that  this  also  is  produced  by  the  action  in  the  heart 
is  perfectly  absurd. 

Some  idea  of  the  force  in  the  arterial  pulsations  may  be  had 
by  watching  the  to-and-fro  movements  in  the  foot  when  the 
legs  are  crossed  at  the  knees  in  the  sitting  posture,  the  foot  of 
the  upper  limb  riding  up  and  down  with  every  rhythm  in  the 
vessels.  To  say  that  the  heart  produces  this  action  also,  is 
utterly  incredible.  Moreover,  this  action  may  be  seen  in  an- 
other form  in  the  fierce  pulsations  which  occur  in  the  local 
vessels  in  inflammatory  processes,  though  the  heart  may 
be  beating  as  usual,  and  which  in  itself  is  ample  proof  of 
automatism  in  the  vessels ;  otherwise  is  inexplicable.  The  pul- 
sations which  occur  in  the  ears  of  a  rabbit  and  wings  of  a  bat 
are  not  synchronous  with  the  action  in  the  heart,  thereby  show- . 
ing  conclusively  independent  action  in  the  vessels  themselves, 
furnishing  incontrovertible  proof  of  this  circumstance. 

Concluding  this  class  of  evidence,  we  may  mention  the  pul- 
sations occurring  in  the  umbilical  cord,  which  are  not  synchro- 
nous with  either  the  maternal  or  foetal  heart,  moreover  occur 
after  connection  is  severed,  and  which  furnishes  absolute  proof 
in  itself  of  automatism  in  the  vessels.  Last,  but  not  least  by  any 
means,  are  the  facts  in  development,  showing  how  a  complex 
circulation  is  carried  on,  with  regular  rhythmical  expansions 
and  contractions  taking  place  in  the  vessels,  in  the  entire 
absence  of  a  heart  for  producing  them;  notably  in  worms. 
The  point  we  wish  to  make  in  this  connection  is,  that  the  ani- 
mal circulation  is  at  first  carried  on  by  the  vessels  only  (inclu- 
sive, of  course,  of  the  actions  in  the  walls  of  the  body  and  the 
molecular  movements),  and  that  the  heart  comes  into  the  vascu- 


148  PHYSIOLOGICAL    ANATOAIY.     . 

lar  system  with  progress  in  development  for  increasing  circu- 
lation simply  ;  being  itself  a  differentiation  in  the  vessels 
for  this  purpose  only,  and  not  to  displace  or  substitute  the 
action  in  the  vessels,  which  can  by  no  means  be  done,  the  local 
actions  themselves  inhibiting  it,  the  question  being  one  of 
increasing  force  in  the  circulation  simply  ;  and  which  is  done 
not  by  the  differentiation  of  this  central  force-pump  only,  but 
also  by  innumerable  muscles  and  nerves  to  the  vessels  them- 
selves;  the  two  being  always  in  correspondence,  the  vessels 
developing  as  the  heart  is  being  developed,  and  growing  pari 
passu  with  each  other.  And  which  holds  true  in  embryogeny 
as  well,  the  nutritive  processes  building  up  the  vessels  contem- 
poraneously with  the  heart,  the  two  going  on  together.  More- 
over, they  are  fundamentally  the  same  in  structure,  consisting 
of  an  inner,  a  middle  and  an  external  layer ;  or  a  serous,  mus- 
cular and  elastic  or  fibrous  tunic  (which  in  the  heart  is 
converted  into  a  pericardial  membrane),  thereby  enabling 
them  to  expand  and  contract  the  same  as  the  heart,  which 
the  functions  call  for.  In  further  proof  of  this  automa- 
tism in  the  vessels,  we  might  mention  also  the  pulsa- 
tions occurring  in  the  post-caval,  axillary  and  iliac  veins  in 
JBatrachia,  which  are  not  synchronous  with  the  action  in  the 
heart ;  also  the  action  in  the  lymph-hearts,  which  are  not  syn- 
chronous with  the  pulsations  in  the  heart ;  nay,  not  even  with 
one  another,  each  seemingly  possessing  an  independent  action 
So,  then,  this  matter  of  automatism  in  the  vessels  rests  upon 
incontrovertible  evidence,  anatomical  and  physiological ;  in 
fact,  everything  is  in  accord  with  this  circumstance,  the  law  in 
the  circulation  also  compelling  it. 

But  the  matter  is  not  yet  fully  summed  up  ;  turn  we  to 
the  special  anatomy  in  the  organs,  to  see  how  this  accords  with 
the  other  phenomena,  or,  as  means  to  ends,  for  producing  the 
work  involved  in  the  tracings,  the  rhythmical  expansions 
and  contractions  in  the  walls  of  the  vessels  indicated  by 
them. 

Descriptive  Anatomy.— 1.  The  arteries  are  very  muscular 
organs,    the    small    arteries    nearly   all    muscles   (Fig.   4G) 


* 


*Tlie  dark  lines  show  the  longitudinal  muscles. 


PHYSIOLOGICAL   ANATOMY. 


l-i9 


If  the  vessels  are  conduits  simply,  what  then  is  the  office  of 
these  numerous  muscles,  seeing  that  muscles  and  nerves  (so 
many  nerves,  too,  to  the  vessels)  are  physiological  adjust- 
ments for  performing  work ;  whether  it  relates  to  circulation  or 
the  voluntary  movements  it  is  all  the  same,  force  being  pro- 


Pig.  46. — Transverse  Section  of  an  Artery  from  a  Vertical  Section  of  the  Skin  of  a 
Guinea  Pig. — Klein,  a,  lumen  of  the  vessel ;  6,  endothelium  seen  in  profile  ;  c, 
intima  ;  d,  circular  muscles  ;  e,  adventitia  ;  /,  cellular  elements  of  adventitia.  (Oc. 
3  ;  obj.  7.) 


Fig.  47. — Longitudinal  Section  of  a  Branch  of  the  Pulmonary  Artery,  from  the  Lung  of 
a  Guinea  Pig. — Klein,  a,  intima  ;  6,  circular  muscular  fibres  cut  across  ;  c,  adven- 
titia,    (Oc.  3  ;  obj.  7.) 


]50  PHYSIOLOGICAL    ANATOMY.    • 

duced  by  this  means,  while  the  energy  is  determined  by  the 
numbers  of  the  muscles  and  nerves  '. 

2.  Why  should  the  muscles  in  the  vessels  be  nearly  all  cir- 
cular, extending  round  the  vessel  '.  (Fig.  47.  b.)  And  since 
muscles  elongate  and  shorten,  or  expand  and  Contract  when  in 
action,  would  not  this  necessarily  expand  and  contract  the 
lumen  correspondingly,  thereby  producing  the  rhythmical 
changes  of  pressure  upon  the  blood  for  speeding  it  through  the 
channels  in  the  manner  as  alleged,  the  vessels  being  coordinated 
with  the  heart  and  the  lungs  by  means  of  the  nerves,  which 
emboss  them  %  It  could  not  be  otherwise,  in  the  very  nature  of 
things,  seeing  that  the  muscles  expand  and  contract,  and 
this  cannot  be  doubted  for  a  single  moment  even.  In  short, 
the  arrangement  of  the  muscles  in  the  vessels  is  the  same  as  in 
the  dermo-muscular  tube  in  the  leech,  which  expands  enor- 
mously (Fig.  17),  only  that  the  longitudinal  muscles  are  more 
numerous  for  effecting  elongation  and  contraction  of  the  body, 
for  which  it  possesses  great  powers  ;  whereas,  in  the  vessels,  the 
object  is  to  effect  expansion  and  contraction  of  the  lumen  for 
producing  the  changes  of  pressure  upon  the  blood  ;  while  the 
less  numerous  longitudinal  muscles  enable  a  degree  of  short- 
ening and  elongation  as  well,  producing  the  so-called  locomo- 
tion in  the  vessels  (Fig.  48),  the  effect  of  elongation  simul- 
taneously with  lateral  expansion,  together  with  the  subsequent 
contraction  which  sets  in,  also  accounting  for  the  retraction  of 
the  vessel  within  the  sheath  after  division,  as  occurs  in  ampu- 
tations, for  example. 

The  movements  connected  with  the  systemic  circulation 
would  include  the  actions  for  producing  a  norm  in  arterial 
pressure,  the  rhythmical  changes  in  pressure  with  inspiration 
and  expiration,  finally  with  the  actions  in  the  heart  itself,  the 
vessels  expanding  during  the  systole  and  contracting  during 
diastole.  But,  in  addition  to  this,  the  local  vessels  expand 
and  contract  for  increasing  and  diminishing  the  lumen,  in 
response  to  special  stimulus  in  the  organs,  whereby  the  local 
circulation  is  increased  or  decreased  correspondingly,  a  matter 
determined  by  the  local  ganglia  or  mind  centres,  which,  as  a 
rule,  are  located  at  the  points  where  the  branches  are  given  off 
from  the  main  stem.    And  the  whole  being  connected  with  the 


PHYSIOLOGICAL    ANATOMY.  151 

medulla  oblongata  by  means  of  the  vaso-motor  centre,  coordi- 
nation with  respiration  and  the  action  in  the  heart  is 
readily  effected ;  at  the  same  time  a  norm  in  pressure  is  main- 
tained. Thus,  these  two  important  anatomical  facts  in  the 
muscles  and  nerves  to  the  vessels  have  easy  and  natural  ex- 
planation 

3.  Last,  but  not  least,  we  have  to  mention  the  progressive 
increase  in  the  numbers  of  the  muscles  in  the  arterial  sys- 
tem from  the  heart  to  the  tissue-territory  (Fig.  49).  How  is 
this  circumstance  to  be  accounted  for  % 

According  to  present  theory,  which  makes  the  heart  the 


Fig.  48. — Elongation  and  Curvature  of  an  Artery  in  Pulsation.— Dalton. 

force  in  the  circulation,  this  arrangement  of  the  muscles  is 
totally  inexplicable,  since  the  effect  would  be  to  place  ob- 
struction in  the  roadway  of  the  blood,  and  hinder  in 
place  of  assisting  the  action  in  the  heart ;  and,  considering 
the  number  of  the  muscles,  it  must  necessarily  involve  pro- 
digious strain  to  the  heart  itself  for  overcoming  the  resistance, 
admitting,  for  the  sake  of  the  argument,  it  were  capable  of  do- 
ing this,  which  can  by  no  means  be  done.  The  vessels  being 
always  full,  arterial  pressure  only  varying,  it  follows  that  in 
order  to  compel  more  blood  into  the  vessels,  this  force  would 
have  to  be  overcome  by  the  heart,  which  is  utterly  impossible. 
Then,  again,  conservation  of  force,  and  of  the  very  blood 
itself,  would  inhibit  such  rude  mechanics,  ignoring  altogether 
strain  to  the  heart,  which,  for  obvious  reasons,  cannot  be 


i52  PHYSTOLOGICAL    ANATOMY. 

done.  And  not  these  only,  but  the  sudden  force  and 
energy  in  the  very  leap  of  the  pulse  would  alike  have  to 
be  explained  by  cardiac  action.  Last,  but  not  least,  the 
danger  to  the  blood  itself  should  it  be  used  as  a  wedge, 
so  to  speak,  to  be  driven  into  the  vessels  by  the  sledge- 
hammer strokes  of  a  central  cardiac  engine  for  forcing 
them  open.  Either  the  delicate  blood-corpuscles  would  go 
to  pieces  in  the  midst  of  this  rude  force,  else  the  aorta 
itself  would  burst,  admitting  the  heart  were  capable  of  per- 
forming the  work  assigned  it,  and  that  the  vast  numbers  of 
circular  muscles  and  elastic  tissue-fibres  could  be  pulled  into 
forcible  extension  by  this  means.  It  is  coarse  and  brutal  to 
the  last  degree,  and  extremely  primitive.  But  the  haemal  me- 
chanics is  of  a  much  higher  order,  nursing  the  blood  and  con- 
serving the  heart  and  vessels  by  reducing  friction  and  resist- 
ance to  a  minimum,  and  which  is  done  by  dividing  the  force 
for  effecting  circulation  into  a  vis  a  f route  and  a  vis  a  tergo, 
acting  in  concert  and  simultaneously,  or  a, pulling  and  a,  push- 
ing force  combined ;  the  former  produced  by  the  arterial  dias- 
tole, the  latter  by  the  cardiac  systole,  working  together  har- 
moniously and  in  conformity  with  the  law  underlying  the* 
organism,  the  blood  flowing  relatively  from  high  to  low 
pressure. 

And  with  the  bases  of  the  arterial  pyramids  in  the  tissues 
and  the  apices  at  the  heart  (Fig.  49),  this  action  in  the  vessels 
would  inevitably  have  that  effect  upon  the  blood  during  the 
cardiac  systole.  And  since  the  pressure  in  the  ventricle  during 
systole  amounts  only  to  about  6  oz.  (mercury),  while  that  in 
the  arteries  falls  below  this,  it  follows  that  during  arterial 
diastole  the  blood  would  flow  from  higl  to  low  pressure, 
pressure  falling  from  the  heart  to  the  tissue-territories  where 
expansion  is  greatest,  being  the  widest  portions  of  the  pyra- 
mids (Fig.  42,  L,  L) ;  and  by  thus  sucking  the  blood,  relieving 
work  at  the  heart  in  proportion ;  while  the  nervous  plexuses 
embossing  the  heart  and  the  vessels  serve  for  coordinating  the 
actions,  as  before  remarked.  And  in  regard  to  these  nerves, 
too,  what  possible  explanation  can  be  found  in  this  old  theory 
for  their  presence  also,  any  more  than  for  the  muscles  ?  None 
whatever,  save  to  produce  contraction  in  the  vessels,  which 


PHYSIOLOGICAL    ANATOMY. 


153 


should  hinder  in  place  of  assisting  the  action  in  the  heart. 
But  they  "relax"  the  vessels  also!  Ah!  that  concedes  the 
matter  fully,  and  no  more  need  be  said,  for  relaxation  means 
expansion  ;  and  the  two  being  coordinated  in  the  functions 
produce  the  phenomena  in  the  tracings.     And  since  expansion 


Fig.  49. — An  Ideal  Diagram  of  the  Arterial  System,  showing  progressive  increase  of  the 
muscles  from  the  heart  to  the  tissue  territory.  L,  L,  upper  and  lower  arterial  pyra- 
mids ;  a,  left  ventricle  and  auricle ;  p,  p,  right  and  left  pulmonary  veins  as  they 
debouch  in  the  auricle  ;  n,  longitudinal  section  of  right  auricle  and  ventricle,  show- 
ing inter- ventricular  and  auricular  septum  ;  B,  B,  walls  of  the  vessels,  representing 
the  progressive  increase  of  the  muscles  from  the  heart  to  the  tissue  territory. 

and  contraction  are  correlated  forces  in  Nature,  the  whole 
mechanics  in  the  animal  circulation  is  at  once  laid  bare  under 
the  white  light  which  is  furnished  in  the  law  underlying  the 


]54  REFLUX    IN  THE   CAPILLAKIES   IMPOSSIBLE. 

organism.  But  should  the  arteries  expand,  would  not  this 
produce  reflux  in  the  capillaries  and  venous  system  \  No  ;  it 
would  not  produce  reflux  in  the  capillaries  and  venous  system, 
for  the  following  unanswerable  reasons  inhibiting  it,  notably  : 
1.  Pressure  in  the  arteries — thanks  (!)  to  the  force  in  the  heart — 
is  ever  higher  than  in  the  capillaries  and  venous  system;  hence  re- 
flux in  the  latter  is  made  impossible.  2.  Expansion  in  the  vessels 
is  from  the  heart  to  the  tissue-territories,  while  the  blood 
keeps  pace  with  it,  flowing  into  the  vessels  as  rapidly  as  ex- 
pansion is  effected ;  and  even  in  excess  of  this,  transmitting 
some  of  the  force  in  the  heart  to  the  walls  during  the  systole, 
thus  producing  an  increase  of  pressure  during  this  time,  which 
is  still  further  augmented  by  the  systole  in  the  vessels,  which 
sets  in  at  the  end  of  the  cardiac  systole,  the  one  running  into  the 
other,  the  same  as  in  the  respiratory  rhythms,  with  which  the 
vessels  are  coordinated,  as  in  the  heart,  the  principle  being  the 
same  ;  while  reflux  in  the  arteries  is  obviated  by  the  valves  at  the 
root  of  the  aorta  and  pulmonary  arteries,  so  that  reflux  in  the 
arterial,  capillary  or  venous  systems  cannot  possibly  occur  ;  the 
only  thing  being  venous  stasis  in  the  systemic  capillaries  from 
damming  of  the  blood  at  the  right  side  of  the  heart  by  obstruct- 
ing circulation  in  the  lungs,  the  venous  system  filling  back  to 
the  capillaries.  And  so  far  as  the  argument  is  concerned,  we 
might  safely  rest  the  case  here,  were  it  not  incumbent  upon  us 
to  carry  the  matter  further  by  explaining  all  the  phenomena, 
anatomical  and  physiological,  appertaining  to  the  vessels  ;  and 
this  a  true  theory  of  the  circulation  should  be  capable  of 
doing,  else  it  is  not  the  true  theory,  but  a  false.  And  by  apply- 
ing this  crucial  test  to  the  Harveian,  its  utter  fallacy  will  at 
once  be  seen,  for  it  would  leave  all  these  matters  unex- 
plained and  inexplicable  ;  and  the  beautiful  adjustments  that 
obtain  in  the  vessels  and  in  the  heart  for  producing  work  in 
the  circulation  would  be  perfectly  meaningless.  But  by  the 
true  theory  it  is  all  brought  out  fully  and  elucidated  so  that 
everything  is  seen  to  be  just  as  it  should  be  to  meet  the  require- 
ments in  the  functions  and  the  ends  sought  to  be  accomplished, 
while  the  enormous  resources  in  Xature  and  her  matchless 
handiwork  receive  additional  emphasis,  inspiring  a  feeling  of 
awe  and  reverence  in  the  mind  unspeakable  ! 


AUTONOMY   IN    THE   VESSELS    INEVITABLE.  155 

Proceeding  with  the  anatomy,  we  come  next  to  the  thick 
elastic   coat  in  the  vessels       The  animal  circulation  being 
based  upon  pressure,  for  increasing  pressure  in  the  arteries 
it  calls  for  this  thick  elastic  coat  for  producing  it,  otherwise  it 
were  impossible  to  effect  it  (as  is  seen  in  the  case  of  the  veins), 
the  vessels  giving  way  under  it ;  and  which  would  also  explain 
the  circumstance  of  the  relative  thickness  of  the  elastic  coat  in 
the  aorta,  together  with  its  progressive  increase  as  the  heart  is 
i  approached,  where  pressure  is  greatest,  the  artery  at  this  point 
being  nearly  all  elastic  fibres.     For  maintaining  high  pressure 
in  the  arteries,  then,  is  the  explanation  of  this  strong  elastic 
coat  in  the  vessels,   the  whole  matter  relating  to  pressure ; 
while  the  high  pressure  which  is  produced  in  the  arteries  by 
the  action  in  the  ventricles  serves  for  increasing  the  capillary 
circulation,  at  the  same  time  that  it  functions  as  the  vis  a 
tergo  to  the  venous  system  for  compelling  this  blood  more 
rapidly  toward  the  lungs,  the  arteries  contracting  and  expand- 
ing synchronous  with  respiration  for  increasing  the  action,  and 
for  compelling  the  blood  more  rapidly  through  the  lungs  dur- 
ing inspiration  and  expiration,  as  before  remarked. 

Concerning  the  Local  Actions. — Finally,  we  come  to  the 
local  actions  in  the  vessels  in  connection  with  the  multitudi- 
nous functions  in  the  body,  and  which  vary  from  moment  to 
moment,  so  that  perfect  uniformity  in  the  circulation  nowhere 
obtains,  and  is  utterly  impossible.  Now,  then,  for  producing 
these  actions,  it  calls  for  automatism  ia  the  vessels  in  order  to 
enable  them  to  respond  to  stimulus  so  as  to  increase  or  decrease 
the  circulation,  as  the  case  may  be.  And  in  the  midst  of  this 
overwhelming  avalanche  of  facts,  what  is  to  become  of  that  old 
monistic  theory  of  cardiac  action  only? — it  is  utterly  lost, 
swallowed  up,  disappearing  from  sight  in  the  midst  of  the 
myriad  phenomena. 

It  need  not  detain  us;  one  or  two  illustrative  cases  will 
serve.  Thus,  when  food  is  passing  into  the  stomach  its  pres- 
ence produces  enormous  afflux  of  arterial  blood  in  the  organ, 
the  vast  capillary  network  in  the  gastric  mucous  membrane 
(Figs.  64,  65)  springing  open  at  once,  for  compelling  blood 
into  the  parts  in  the  measure  of  the  physiological  require- 
ments in  the  glandular  structures  and  for  increasing  absorp- 


156  AUTONOMY   IN    THE   VESSELS    INEVITABLE. 

tion,  at  the  same  time  the  biliary  and  pancreatic  secretions 
are  increased  correspondingly,  which  is  done  by  springing  open 
the  vascular  feeders  in  the  separate  arteries  of  the  coeliac  axis, 
while  this  in  turn  has  regulation  by  the  amount  of  food  ingested, 
diminishing  with  the  gradual  absorption  and  removal  of  the  gas- 
tric contents.  And  each  organ,  by  having  a  separate  feeder  (Fig. 
110),  controlled  by  the  local  nerves  and  ganglia  (Fig.  109),  is 
thus  enabled  to  command  all  the  blood  that  is  needed  for  effect- 
ing the  functional  processes  without  interfering  with  the  general 
circulation,  which  must  not  be  done.  K  ay,  more  than  this,  each 
group  of  cells  is  enabled,  by  the  separate  branches  given  off 
by  the  common  feeder  to  the  organ,  to  command  the  blood 
they  need  in  the  special  exigencies  arising  in  the  local  actions, 
so  that  the  organ  as  a  whole  and  in  its  several  parts  may  com- 
mand the  blood-suppiy  in  the  measure  of  the  requirements. 
And  so,  likewise,  down  the  whole  intestinal  canal,  each  and 
every  portion,  by  means  of  the  separate  feeders  to  them  repre- 
sented in  the  trunks  and  numerous  branches  of  the  superior 
and  inferior  mesenteric  arteries  (Figs.  Ill,  112),  may  com- 
mand, by  means  of  the  special  nerves  and  ganglia,  the  amount 
of  blood  they  need  without  interfering  with  any  other  por- 
tion, so  that  perfect  independence  of  action  is  maintained. 
Well,  while  all  this  is  going  on  in  digestion,  the  heart  beats 
as  usual,  and  as  though  there  were  no  such  thing  as  digestive 
processes,  and  all  the  multitudinous  arrangements  for  expe- 
diting and  controlling  the  flow  of  blood  into  the  organs : 
moreover,  is  utterly  incapable  of  effecting  the  actions,  requiring 
automatism  in  the  vessels  themselves  in  order  to  accomplish  it. 
But  the  heart,  by  maintaining  pressure  in  the  arterial  system, 
which  tends  to  fall  by  reason  of  these  depletions,  does  subserve 
important  uses,  the  vaso-motor  system  also  contributing  by  con- 
tracting them  in  proportion,  and  thus  maintaining  pressure, 
which  is  essential  for  increasing  the  local  actions  by  causing 
instantaneous  flow  into  the  capillaries  the  moment  they  ex- 
pand, at  the  same  time  maintaining  the  other  actions  in  the 
body  which  the  scheme  calls  for. 

Take  another  case — the  kidneys,  for  example.  Upon  occa- 
sion an  enormous  flow  of  urine  takes  place.  Well,  since  this 
is  drawn  directly  from  the  blood  through  the  secretory  actions 


NERVOUS   SUPPLY   TO   THE   VESSELS. 


157 


in  the  organs,  it  follows  that  there  must  be  an  increased  supply 
of  blood  in  the  parts  in  order  to  effect  it.  And  how  is  it  pos- 
sible to  command  this  increased  supply  of  blood  save  by  spring- 
ing open  the  renal  vessels  correspondingly  ?  It  all  comes  from 
the  blood,  and  there  must  be  increase  of  this  in  the  organs  for 
producing  it,  the  amount  being  determined  by  lumen  of  the 
vessels,  which  the  local  nervous  ganglia  regulate  by  means  of 
the  nerves  extending  over  and  literally  embossing  them  (Fig. 
109,  9,  G).  At  the  same  time,  the  vessels,  arteries  and  veins  are 
very  muscular  And  there  is  no  nervous  plexus  in  the  body 
richer  in  nerves  and  nervous  ganglia  than  the  renal,  save  the 
solar  plexus  itself,  the  solar  centre  of  the  intestinal  mechanics, 


Fig.  50. — Ramification  of  Nerves  and  Termination  in  the  Muscular  Tunic  of  a  Small 
Artery  of  a  Frog. — Arnold. 

of  which  it  forms  a  part,  being  immediately  contiguous  to  it 
(Fig.  109,  G) ;  thereby  showing  the  relative  importance  of  the 
renal  functions  for  maintaining  a  balance  in  the  organism 
through  the  excretory  functions  in  these  organs.  But  in  the 
absence  of  the  power  to  expand  and  contract  the  vessels,  all 
this  would  be  meaningless.  As  it  is,  everything  is  in  corre- 
spondence for  promoting  the  ends  in  view. 

Nervous  Supply  to  the  Vessels. — Finally,  we  come  to  the  nerv- 
ous supply  to  the  vessels,  with  the  mode  of  termination  in  the 
sarcous  elements  for  effecting  the  actions  spoken  of.   According 


]58  NEKV0US   SUPPLY    TO   THE   VESSELS. 

to  the  exhaustive  researches  of  Arnold,  the  nervous  twigs  in  un- 
striped  muscles  consist  partly  of  medullated  and  partly  of  non- 
medullated  fibres,  in  varying  proportions,  penetrating  to  the 
nucleus  of  the  muscle  cell  in  the  form  of  a  fine  terminal  fila- 
ment, and  ending  probably  in  the  nucleolus.  Externally,  in 
the  connective-tissue  covering  the  muscle,  these  nerves  are 
arranged  in  the  form  of  a  wide-meshed  network,  in  which,  as 
Beale  has  pointed  out,  ganglion  cells  are  to  be  found  at  cer- 
tain points  in  the  muscles  of  the  vascular  system  (Frey).  To 
this  the  name  ' '  ground  plexus' '  has  been  given.  From  this  are 
given  off  the  nerves  which  penetrate  between  the  layers  of 
muscles  to  form  the  ' '  intermediate  network' '  (Fig.  50),  which 
lies  immediately  upon  or  between  the  muscular  layers  ;  while 
from  this  again  a  series  of  fibres  are  given  off  which  penetrate 
between  the  muscle  fibres  to  form  a  new  network,  "intra- 
muscular," surrounding  the  muscle  cells,  from  which  dark 
straight  fibres  of  extreme  fineness  penetrate  into  the  Cells,  and, 
advancing  to  the  nucleus,  terminate  in  the  nucleolus  (Franken- 
hauser).  The  number  of  terminal  filaments  which  enter  any 
one  muscle  cell  corresponds  with  the  number  of  granules 
occurring  in  the  nucleus  (Frey).  According  to  Arnold,  these 
fibrillse  leave  the  nucleoli  again  in  the  opposite  direction,  and 
after  having  traversed  the  nucleus  and  body  of  the  cell,  unite 
once  more  with  the  intra-muscular  network.  Finally,  a  set  of 
pale  fibres  proceed  directly  from  the  "ground  plexus"  to  the 
muscle  cells,  forming  a  direct  connection  between  them.  Now, 
then,  what  can  be  the  purpose  of  the  ganglion  cells  in  the 
"ground  plexus"  and  the  intricate  nervous  arrangements 
which  obtain  in  arterial  muscles  for  producing  molecular 
action  in  the  cells  other  than  to  effect  the  actions  spoken  of  \ 
Seeing  also  that  an  intimate  connection  subsists  between  them 
and  the  plexuses  embossing  the  vessels,  connecting  them  with 
the  central  nervous  system.  It  is  needless  to  extend  the  matter. 
This  brings  us  to  the  remaining  element  in  the  arterial  tracings, 
notably,  dicrotism ;  but  that  will  require  a  chapter  to  itself, 
involving  the  action  in  the  capillaries  in  connection  with  res- 
piration in  the  tissues. 


CHAPTER  VIII. 

RESPIRATION     IN     THE     TISSUES     AND     THE     ACTION      OF     THE 
CAPILL ABIES    IN    CONNECTION    THEREWITH. 

Two  Respiratory  Movements  Going  on  at  the  Same  Time  in  the  Body,  one  in  the  Lungs, 
the  other  in  the  Tissues — The  Force  for  Producing  the  Movements  Propagated  from 
the  Medulla  Oblongata  and  Respiratory  Centre — The  Composite  Character  in  Arterial 
Tracings  Readily  Explained — Rhythmical  Expansions  and  Contractions  in  the  Tissues 
Synchronous  with  the  Actions  in  the  Lungs,  Heart,  Arteries  and  Capillaries — The 
Waves  Superposed  Upon  One  Another  in  the  Order  Named,  or  the  Cardo- Arterial 
Upon  the  Respiratory,  and  the  Capillary  or  Dicrotic  Waves  Upon  the  Cardo- 
Arterial — Capillary  Action  the  Source  of  Dicrotism — Mode  of  Demonstrating 
this  Circumstance — The  Action  in  the  Capillaries  Producing  a  Current  Into  and 
Out  of  the  Tissue-Interstices — Relations  of  the  Cell-Brood  to  this  Circulation — 
Nervous  Apparatus  for  Connecting  Them  with  the  Capillaries  and  Central  Ner- 
vous System  for  Increasing  and  Diminishing  the  Local  Circulation,  with  the  Exi- 
gencies in  the  Functions — Mechanics  in  Blushing — Explanation  for  Arrest  of  Arterial 
Pulsations  Upon  the  Distal  Side  in  Aneurisms — Action  in  the  Venous  System — 
Functions  of  the  Muscles  and  Nerves  in  Veins — The  Relations  they  Sustain  to 
Respiration — Explanation  for  the  Great  Volume  of  Venous  Blood  and  the  Slowness 
in  this  Circulation — Also  for  the  Motion  in  the  Brain  Synchronous  with  Respiration. 

Finally,  in  order  to  fully  interpret  the  phenomena  in  arterial 
tracings,  it  will  be  necessary  to  consider  them  in  connection 
with  the  capillary  circulation  and  the  respiration  in  the  tissues. 

In,  short,  there  are  two  respiratory  movements  going  on 
at  the  same  time  in  the  body :  One  in  the  lungs,  the  other 
in  the  tissues,  and  which  answer  to  the  two  poles  in  the 
floral  circulation;  while  the  heart  and  vessels  function  as  a 
carrier  between  them,  expanding  and  contracting  upon  the 
fluids  for  increasing  circulation  in  the  two  poles,  while  the 
force  for  compelling  these  actions  is  transmitted  over  the 
vessels  from  the  central  nervous  system  by  means  of  the 
nervous  plexuses  which  emboss  them.  The  result  is  the  pro- 
duction of  the  composite  character  in  the  arterial  tracings,  or 
the  existence  of  respiratory,  cardo -arterial  and  dicrotic  waves  ; 
and  which  are  superposed  one  upon  the  other  in  the  order 
named,  or  the  respiratory  by  the  cardo-arterial,  and  the  latter 


160  RESPIRATION   IN   THE  TISSUES. 

by  the  capillary  or  dicrotic  waves,  which  represent  the  respi- 
ratory action  taking  place  in  the  tissues.  It  is  all  plain 
enough  and  readily  understood,  the  actions  being  simply 
throbbed  over  the  vessels  as  the  common  carrier  between  the 
two  poles  of  the  circulation  from  the  medulla  oblongata,  in 
which  the  nerves  converge  for  coordinating  and  unifying  the 
actions  throughout,  which  the  scheme  calls  for. 

This  being  the  case,  of  course  there  should  be  a  means  of 
proving  and  demonstrating  the  fact  by  showing  the  following 
circumstances : 

First,  That  the  respiratory  waves,  or  the  waves  produced  by 
respiration  in  the  lungs,  reach  and  pervade  the  tissue-terri- 
tory. 

Second,  That  the  cardo-arterial  loaves,  or  pulsations,  reach 
and  pervade  the  tissue-territory. 

Third,  That  dicrotism,  so-called,  is  the  capillary  movement 
superposed  upon  the  cardo-arterial. 

Thanks  (!)  to  the  distinguished  physiologist  and  experimen- 
talist at  Paris,  we  find  the  evidence  ready  to  our  hand :  It 
consists  in  inclosing  the  hand  in  a  glass  jar  filled  with  water, 
made  air-tight  over  the  top,  and  so  constructed  as  to  cause  the 
oscillations  it  produces  in  the  water  to  be  transmitted  to  the 
lever  of  a  sphygmograph  for  registering  the  result  (Pig.  51).* 
It  produces  the  following  tracings  (Fig.  52,  V) :  f 

In  the  same  cut  the  tracings  are  contrasted  with  the  tracings 
produced  by  the  arteries  (C),  but  for  a  different  purpose  from 
the  present,  the  author  being  under  the  impression  that  they 
are  produced  by  the  action  in  the  heart,  alleging  the  same 
for  the  arterial  tracings,  making  this  organ  the  force  in  the 
circulation,  which  cannot  be  thought  of  for  a  single  moment, 
for  the  reasons  already  given. 

We  will  begin  with  respiration,  as  this  is  the  basis,  so  to 
speak,  of  animal  life,  beiag  the  pumping  action  for  compelling 
the  commerce  in  the  organism  with  which  the  vascular  system 
connects  for  distributing  it  through  the  tissues,  as  before  re- 

*  La  Methode  Graphique  dans  les  Sciences  Experimental,  et  Principale- 
ment  en  Pbysiologie  et  en  Medecine.  Par  E.  J.  Marey,  Professeur  au  College 
de  France,  Membre  de  l'Acadernie  de  Medecine.     Fig.  327. 

t  Ibid      Fig.  328. 


RESPIRATION   IN   '1HK   TISSUES. 


161 


marked.     Now,  then,  we  have  this  great  movement,   which 
pervades  the  entire  organism  from  centre  to  circumference, 


Fig.  51.— Apparatus  for  Inscribing  the  Changes  of  Volume  in  the  Hand.  The  membrane 
which  traverses  the  fluid  is  rendered  immovable  by  a  metallic  plate  ;  while  the 
oscillations  which  are  produced  in  the  water  are  transmitted  through  the  bulb  and 
vertical  tube  to  the  lever  of  the  sphygmograph  by  means  of  air- — Marey. 


Fig.  53.— Sphygmographic  Tracings,  showing  the  undulations  in  the  vessels  and  tissues 
synchronous  with  respiration.  C,  respiratory  rhythms  in  the  arteries  ;  A,  crest  of  a 
wave  ;  C,  trough  of  a  wave.  V,  respiratory  rhythms  in  the  tissues,  smaller,  but 
very  apparent  ;  b,  trough  of  a  respiratory  wave  in  the  tissues,  corresponding  with 
the  crest  (A)  to  the  respiratory  wave  in  the  arteries,  which  arrives  later  ;  £>,  crest  of 
a  respiratory  wave  in  the  tissues,  which  corresponds  with  the  trough  (C)  of  the 
respiratory  wave  in  the  arteries  which  had  passed  into  the  tissues. 


1G2  RESPIRATION   IN   THE  TISSUES. 

registered  in  the  arterial  tracings  in  the  large  undulations 
which  occur  in  it  (Fig.  52,  C,  A).  Two  and  one-half  such 
waves  occur  in  the  tracings,  the  one  at  the  right-hand 
side  of  the  figure  being  cut  off.  And  the  same  circumstance 
also  presents  in  the  tissue  tracings  (V),  only  that  they  are 
smaller  and  arrive  later,  having  to  pass  over  the  arteries  in 
order  to  reach  the  tissues,  the  nerves  lying  upon  the  walls  of 
the  vessels.  As  a  result  of  this  circumstance,  therefore,  the 
crest  (A)  of  the  respiratory  waves  in  the  arteries  corresponds 
with  the  trough  (b)  of  the  respiratory  waves  in  the  tissues,  and 
vice  versa  (C,  D).  In  other  words,  the  same  number  of  res- 
piratory actions  occur  in  the  tissues  as  in  the  arteries  to  the 
very  fractional  portions  of  the  waves,  and  nothing  could  be 
more  conclusive  of  respiratory  action  in  the  tissues  than  this 
very  circumstance.  Prof.  Marey,  however,  conceiving  it  to  be 
due  to  the  action  in  the  heart,  has  drawn  perpendicular  lines 
at  certain  points  through  the  upper  and  lower  tracings,  in 
order  to  show  correspondence,  as  he  thinks,  between  the  action 
in  ih  e  heart  and  the  changes  of  volume  in  the  hand,  as  cause 
and  effect.  But  we  fail  to  perceive  it  in  this  light,  for  the  rea- 
sons already  given.  It  is  not  so  simple  as  this,  but  highly 
complex,  the  result  of  several  actions  blended  together.  It 
will  not  answer  at  all. 

What  is  to  be  done  with  the  correspondence  between  the 
large  undulations — the  two  whole  and  the  fractional  portions 
of  the  respiratory  waves — which  occur  in  both  tracings,  since 
there  is  exact  correspondence  ;  only  they  are  smaller  in  the 
tissues,  as  a  matter  of  course,  and  arrive  later,  which  is  also  a 
matter  of  course.  Then  there  are  sixteen  smaller  waves,  which 
are  superposed  upon  the  larger,  indenting  them  like  the  teeth 
in  a  saw,  which  answer  to  the  action  in  the  heart  and  arteries, 
the  same  number  precisely  occurring  in  both  tracings,  so  that 
correspondence  here  is  also  complete — impossible  to  make  it 
more  so.  Well,  what  is  to  be  done  with  this  correspondence  % 
as  it  all  must  be  explained  and  satisfactorily  accounted  for  ;  else 
the  theory  is  false.  These  smaller  undulations  occur  all  along 
the  surface  of  the  respiratory  waves,  at  the  crest  (A),  at  the 
sides,  and  in  the  very  trough  (C)  itself,  in  regular  order  and 
succession ;    the    same    applying  for  both  tracings.     Then, 


AKKESTING  DICROTISM.  163 

again,  there  are  four  of  these  to  each  of  the  billows,  which 
correspond  with  the  relative  frequency  of  the  respiratory 
and  cardiac  rhythms,  showing  conclusively  that  the  one  is 
respiratory,  the  other  cardiac,  or,  rather,  cardo-arterial,  being 
produced  by  the  united  actions  of  the  heart  and  arteries.  The 
action  in  the  heart,  forsooth  !  There  is  more  here  than  is  ex- 
plicable by  the  action  in  the  heart.  It  cannot  produce  the  res- 
piratory undulations  in  the  arteries,  and  why  should  it  pro- 
duce the  corresponding  undulations  in  the  tissues  %  It  follows 
that  the  respiratory  waves,  as  well  as  the  waves  produced  by 
cardo-arterial  movement,  pervade  the  tissues,  the  latter  being 
superposed,  so  to  speak,  upon  the  former,  giving  rise  to  the 
serrated  appearance  in  the  respiratory  waves.  But  we  have 
still  smaller  waves,  which  in  turn  are  superposed  upon  the 
arterial,  producing  the  appearance  known  as  dicrotism,  and 
indenting  these  as  the  arterial  the  respiratory,  and  which  also 
present  in  both  the  tracings,  bat  are  more  distinct  in  the  lower 
( V) ;  and  which,  also,  must  be  accounted  for  with  the  others. 
Concerning  Dicrotism. — The  very  fact  that  dicrotism  is 
more  pronounced  in  the  tissue  tracings  (Fig.  52,  v)  is  of  itself 
suggestive  of  capillary  action  as  its  source,  and  that  it  is  ac- 
tually produced  by  the  action  in  the  capillaries  is  also  easily 
proven  by  simply  arresting  the  action  in  these  vessels  when 
dicrotism  promptly  disappears,  but  re-appearing  again  with 
restoration  of  capillary  action,  at  once  showing  the  relations 
they  sustain  to  each  other.  This  object  is  attained  by  dis- 
tending the  vessels  with  venous  blood,  and  so  preventing 
their  action  ;  while  the  action  is  reestablished  again  by  reliev- 
ing them  of  the  blood.  Thus,  with  the  hand  in  the  registering 
apparatus  as  previously  (Fig.  5] ),  a  constricting  band  was  placed 
above  the  elbow  sufficient  to  arrest  the  venous  blood  only, 
and  after  waiting  a  short  interval  for  the  blood  to  accumulate 
in  the  hand,  was  again  relaxed.  When  suddenly  and  forcibly 
applied  and  as  suddenly  relaxed,  it  produced  the  upper  trac- 
ing (Fig.  53)  ;*  when  slowly  applied  and  slowly  relaxed,  the 
lower. 

*  (Ibid,  Fig.  330.)  It  is  proper  to  remark,  in  this  connection,  that  Professor 
Marey  did  not  have  the  present  purpose  in  his  mind  when  he  made  this  experi- 
ment, but  we  thank  him  all  the  same  for  the  important  contribution  to  science. 


16-i  ARRESTING  DICROTISM 

It  will  be  seen  by  the  upper  tracing  that  when  the  venous 
blood  is  entirely  arrested,  dicrotism  promptly  disappears ; 
nevertheless,  the  arterial  movements  corresponding  with  the 
action  in  the  heart  are  well  pronounced,  dicrotism  only  disap- 
pearing ;  doing  so  almost  at  once  after  the  veins  are  constricted. 
And  this  shows  that  it  is  due  to  the  choking  of  the  capillaries 
by  the  damming  of  venous  blood  in  them,  preventing  their 
action.  At  least,  this  would  account  for  the  circumstance. 
This  is  also  corroborated  by  the  second  tracing,  in  which 
the  blood  was  not  entirely  arrested,  but  nearly  so  toward  the 
middle  or  summit  of  the  tracing,  where  dicrotism  disappears 
momentarily  only,  showing  some  action  still  left  in  the  capil- 
laries. In  other  words,  the  rhythms  in  the  large  vessels  con- 
tinue and  are  well  pronounced,  but  dicrotism  disappears.  The 
sudden,  almost  perpendicular,  fall  in  the  upper  curve,  which 
was  due  to  the  energetic  contraction  in  the  capillaries  and  veins, 
produced  by  the  venosity  of  the  blood,  which  acts  as  a  stimulus, 
will  give  some  idea  of  the  force  in  these  vessels,  which  should 
not  be  lost  sight  of  in  the  mechanics  of  circulation,  at  once  show- 
ing that  the  force  is  available  in  the  circulation  by  coordinating 
the  capillaries  and  veins  with  respiration,  as  dcfes  actually 
occur ;  moreover,  is  fully  proven  by  the  tissue-tracings  (Fig. 
52,  V).  In  other  words,  the  same  principle  precisely  obtains 
for  increasing  circulation  in  the  capillaries  and  veins  as  in  the 
lungs,  heart  and  arteries — namely,  by  rhythmical  changes  in 
pressure ;  while  the  force  is  propagated  from  the  medulla 
oblongata  by  means  of  the  nerves,  connecting  them  with  the 
respiratory  centre  (Fig.  54),*  the  same  as  in  the  other  cases  ; 
since  this  is  necessary  for  producing  continuity  in  force  for 
maintaining  an  uninterrupted  current  of  the  blood  in  the  round 
of  the  circulation,  in  the  measure  of  the  physiological  require- 
ments, otherwise  impossible.     It  is  plain  enough. 

The  damming  of  the  venous  blood  in  the  capillaries  by  dis- 
tending them  must  inevitably  choke  the  pumping  action,  the 
blood  being  unable  to  escape  by  reason  of  the  ligature,  while 
the  energy  with  which  they  contract  to  force  out  the  blood  is 
seen  in  the  sudden  fall  of  the  upper  curve  the  moment  the 
ligature  is  relaxed. 

*  Handbook  for  the  Physiological  Laboratory,  plate  xxxvi.  Burd  on -Sander- 
sou,  etc. 


ARRESTING-  DICE0TI5M. 


165 


3 

err-  • 

§    « 

cr  o 
©   ^ 

II' 


CO 


"1    «. 

M2 


3  3 


3 

a" 

rt 

r* 

a 
-x 

el- 

CO 

d 

s' 

B  da 
*2  2 
St1  ^ 


1    o 

<*■  o 


CD     ^ 

8  ■*- 
o  5* 
e  ° 

e+    O 

J-h  CD 
I" 

Is 

d  p- 


P    1 


§1 

d-  >-t 

cd   p 
d    o 

CD     B 


C?    » 

&  d- 

*    >o 

»d 


166  DICROTISM   DUE   TO   CAPILLARY   ACTION. 

When  gradually  relaxed,  the  escape  of  blood  is,  of  course, 
in  correspondence  ;  hence  the  difference  in  the  curves  of  the 
two  tracings.  It  is  not  produced  by  the  force  in  the  heart, 
as  the  eminent  Frenchman  conceives  ;  but  the  force  is  upon 
the  ground  where  the  work  is  done,  and  in  the  very  walls  of 
the  vessels  themselves,  which  regulate  their  own  circulation. 
The  heart  is  too  far  removed  for  this  work,  were  not  the 
principle  itself  wrong.     The  pith  of  the  whole  matter  being 


Fig.  54.— Horizontal  Section  of  Tongue  of  Frog  Treated  with  Chloride  of  Gold,  showing 
the  distribution  of  non-medullated  nerve  fibres  to  a  capillary  blood-vessel,  a,  capil- 
lary vessel  ;  b,  coarse  non-medullated  nerve  fibres  ;  c  and  d,  fine  non-medullated 
nerve  fibres  forming  a  plexus  which  surrounds  the  vessel  like  a  sheath  ;  d,  non- 
medullated  nerve  fibres  in  the  wall  of  the  vessel.     (Oc,  3  ;  obj.,  8.)— Klein. 

that  the  tissues  respire  for  maintaining  the  life  that  is  in 
them  and  for  increasing  their  activities,  and  they  must  con- 
nect with  the  respiratory  centre  and  the  action  in  the  lungs 
in  order  to  command  the  oxygen  that  is  needed  for  increasing 
metabolism  in  the  measure  of  the  requirements,  respiration  in 
consequence  readily  rising  and  falling  with  the  swell  in  the 
activities.  And  the  force  for  effecting  this  action  in  the  capil- 
laries being  propagated  over  the  vascular  lines  from  the  me- 
dulla oblongata  would  easily  account  for  dicrotism  in  the 


DICROTISM   DUE  TO   CAPILLARY   ACTION.  167 

heart  and  arterial  system  ;  and  why  this  should  be  more  pro- 
nounced with  the  increase  of  venosity  in  the  blood,  for  this 
makes  the  demands  in  the  cell-brood  more  imperious,  at  the 
same  time  it  acts  as  a  stimulus  to  the  nervous  centres.  It 
would  not  do,  then,  to  have  this  important  link  in  the  vas- 
cular chain  neglected,  as  it  would  involve  failure  to  all  the 
rest.  In  fine,  the  large  undulations  which  answer  to  the  pump- 
ing action  in  the  lungs  are  let  down,  as  it  were,  in  stages  till  it 
meets  the  finer  actions  in  the  capillaries,  and  so  passing  on 
into  the  venous  system,  while  the  whole  is  intimately  con- 
nected by  means  of  the  nerves  extending  over  them  for  pro- 
ducing an  uninterrupted  flow  of  blood  between  the  lungs  and 
cell-brood  (the  two  poles  in  the  circulation),  the  cells  respir- 
ing through  the  lungs,  as  it  were,  by  means  of  the  arteries  and 
veins  which  bring  in  the  oxygen  and  carry  out  carbonic  acid, 
or  upon  the  same  principle  precisely  as  obtains  in  the  embryo, 
which  breathes  through  the  placenta  by  means  of  the  um- 
bilical arteries  and  vein,  only  that  in  the  latter  there  is  still 
greater  removal  from  the  environment  or  source  of  supplies. 
Still,  the  principle  is  the  same  in  both,  and  the  mechanics 
must  connect  through  and  through  in  order  to  accomplish  it. 
It  would  require  that  all  the  parts  should  be  coordinated  in  the 
vascular  chain  in  order  to  produce  and  unify  the  universal 
pumping  actions  for  increasing  circulation  commensurate  with 
the  physiological  requirements,  and  dicrotism  must  take  its 
place  in  the  march  of  the  phenomena,  at  the  same  time  it 
shows  the  interdependence  subsisting  between  them  and  the 
common  relation  they  sustain  to  the  law  underlying  the  organ- 
ism, the  whole  performing  in  harmonious  concert  under  action 
of  the  nervous  forces  radiating  from  the  solar  centre  of  the 
organism.  So,  then,  we  can  readily  understand  why  the  capil- 
lary or  dicrotic  waves  should  announce  themselves  in  the  tissue 
tracings,  appearing  and  disappearing  with  the  relaxing  and 
tightening  of  the  ligature  which  relieves  or  chokes  the  capil- 
laries, as  the  case  may  be.  In  this  manner,  then,  that  vexed 
problem  in  experimental  physiology  has  easy  and  natural 
solution,  dicrotism  falling  into  line  at  the  proper  time  and 
place  with  the  other  phenomena,  leaving  nothing  to  explain, 
which  a  true  theory  of  the  circulation  would  naturally  do. 


118  DICROTISM   DUE  TO   CAPILLARS   ACTION. 

Another  Mode  of  Proving  Capillary  Action. — Another 
mode,  the  very  opposite  of  this,  obtains  for  proving  the  action 
in  the  capillaries  as  a  potential  and  independent  factor  in  the 
circulation,  without  which  neither  the  systemic  nor  local  cir- 
culation could  be  made  commensurate  with  the  physiological 
requirements,  and  animal  life  would  have  arrest  at  the  flora. 
It  consists  in  arresting  the  flow  of  blood  in  the  main  arterial 
trunk  or  feeder  for  the  limb,  but  leaving  the  veins  open  ;  and 
by  thus  cutting  off  the  force  in  the  left  ventricle,  to  throw  the 
burden  of  the  tissue  circulation  upon  the  local  vessels  them- 
selves, which  should  give  us  some  idea  of  the  force  in  the  local 
vessels  for  compelling  circulation.  The  same  instrument 
answers  for  making  the  tissue  tracings  as  in  the  preceding, 
there  being  no  other  way  of  doing  it,  while  the  evidence  it 
furnishes  is  equally  conclusive.  Of  course,  compressing  the 
main  artery  to  the  limb  at  once  puts  an  end  to  all  the  elements 
of  rhythmic  movement  in  the  tracings,  the  cardo-arterial  with 
the  rest  (Fig.  55,  C,  CY/),  the  uninterrupted  flow  of  blood  in  the 
vessels  being  essential  for  their  development.  The  point  we 
wish  to  make,  however,  is  that  the  hand  immediately  shrinks  in 
volume,  which  the  lever  promptly  registers  in  the  low  depres- 
sion (C,  C)  in  the  curve  of  the  tissue  tracings,  and  this  is 
produced  by  contraction  in  the  vessels,  driving  the  blood 
toward  the  heart  and  lungs ;  and  which  proves  incontro- 
vertibly  an  independent  action  in  the  vessels  themselves.  iSo, 
then,  the  force  to  the  tissue  circulation  is  not  in  the  heart, 
which  singly  assists  it,  the  main  force  being  actually  upon 
the  ground  where  the  work  is  done,  in  the  walls  of  the  vessels 
themselves,  which  regulate  their  own  circulation,  and  must 
do  so  in  order  to  maintain  autonomy  in  the  tissues,  which  is 
essential  to  existence. 

The  sudden,  perpendicular  leap  made  in  the  tissue  tracings 
(a')  when  arterial  compression  is  relieved,  is  the  product  of  the 
combined  action  in  the  vessels,  aided,  of  course,  by  the  press- 
ure in  the  arterial  system,  compelling  the  blood  into  the 
vessels  with  the  energy  of  expansile  action  ;  and  which  soon 
restores  the  normal  rhythm  in  the  vessels  and  the  volume  of 
the  hand,  indicated  by  the  rise  in  the  curve  of  the  tissue  trac- 
ings, sending  it  above  the  horizontal  line,  and  the  reappear- 


THE    FOliCii   IN   THE  TISSUES. 


169 


170  THE   FORCE   IN   THE  TISSUES. 

ance  of  dicrotism  and  arterial  movement.  And  that  it  is 
produced  by  the  action  in  the  vessels  in  the  manner  as  stated 
is  fully  proven  by  the  straight,  perpendicular  line  (c'), 
which  answers  to  diastole  in  the  vessels,  and  which  we 
have  seen  the  heart,  of  itself,  is  utterly  incapable  of  pro- 
ducing unassisted  by  the  action  in  the  vessels,  requiring 
expansile  action  in  the  vessels  themselves  in  order  to  effect 
it.  At  the  same  time,  the  over-distension  which  this  pro- 
duces in  the  capillaries,  the  blood  rushing  into  them  from 
the  pressure  in  the  arterial  system,  tends  momentarily  to 
choke  the  action ;  hence,  the  small  and  imperfect  rhythms 
immediately  succeeding  the  diastole.  Furthermore,  we  know 
that  contraction  must  possess  its  equivalent  in  expansion,  so 
that  the  previous  contraction  in  the  vessels  must  have  its 
representation  in  this  expansile  effort,  the  one  involving  the 
other,  expansion  and  contraction  being  correlated  forces. 
Moreover,  we  have  the  same  circumstance  illustrated  in  blush- 
ing, in  which  there  had  been  no  previous  contraction  from 
anaemia,  the  vessels  suddenly  expanding  under  the  nerv- 
ous force  pouring  into  them  from  the  central  nervous  system, 
the  blood  rushing  into  them  instantaneously  under  the  press- 
ure in  the  arterial  system,  producing  the  characteristic  color- 
ing of  the  skin,  followed  promptly  by  energetic  contraction, 
producing  the  succeeding  blanching  of  the  parts  from  the 
anaemia  this  produces.  So,  then,  there  can  be  no  doubt  what- 
ever of  an  independent  action  in  the  capillaries ;  otherwise, 
these  actions  would  be  meaningless.  Finally,  we  have  to  note 
the  low,  flat,  faint  undulations,  which  answer  to  the  rhythm 
in  the  arteries,  still  remaining  in  the  tissue  tracings,  notwith- 
standing the  occlusion  of  the  brachial  artery  above  the 
elbow,  and  which  marks  the  effort  in  the  vessels  to  perform 
the  normal  functions  inherent  in  them.  But  in  the  absence 
of  the  blood  for  filling  them  pari  passu  with  expansion,  as  in 
the  case  of  the  air  when  excluded  from  the  lungs,  of  course 
they  cannot  expand,  showing  the  effort  to  do  so  only  in  these 
indistinct  and  faint  undulations.  But  this  also  subserves  im- 
portant uses  by  promoting  the  collateral  circulation  in  the 
anastomosing  vessels,  producing  a  suction-force  upon  the 
blood,  the  connecting  capillaries  expanding  under  the  nerv- 


INSPIRATORY    ACTION.  171 

ous  force  pouring  into  them,  and  the  pressure  in  the  ar- 
terial system,  which  rapidly  restores  the  normal  circulation — 
the  one,  the  vis  a  f  route,  the  other,  the  ms  a  tergo,  acting  simul- 
taneously. In  fine,  it  is  the  tremendous  effort  which  the 
tissues  make  to  respire  when  the  supplies  are  cut  off,  threaten- 
ing asphyxia,  and  in  which  everything  is  brought  to  bear  for 
expanding  and  filling  the  vessels  with  arterial  blood. 

Of  course,  any  circumstance  which  should  prevent  the  flow 
of  blood  through  the  arteries  would  also  be  reflected  in  the 
tissue  tracings,  since  this  would  inhibit  afflux  of  blood, 
which  is  essential  for  filling  the  vessels  pari  passu  with 
expansioD,  in  order  to  effect  expansion  ;  otherwise  impossible. 
In  other  words,  the  capillaries  cannot  expand  unless  the  blood 
flows  into  them  simultaneously,  since  it  would  require  pro- 
digious force  to  effect  it  in  the  absence  of  the  blood,  involv- 
ing the  production  of  a  vacuum  throughout  the  whole  affected 
capillary  territory,  and  which  the  capillaries  are  by  no 
means  capable  of ;  hence,  in  this  condition  they  contract  till 
the  lumen  is  entirely  closed  (if  the  interruption  is  com- 
plete), and  the  parts  are  blanched.  But  with  the  blood 
flowing  into  them  all  the  while,  as  in  the  normal  condition, 
there  is  no  difficulty  in  effecting  expansion,  while  the  high 
pressure  in  the  arterial  system  in  warm-blooded  animals 
enables  it  to  be  made  very  energetic,  and  which  the  exigencies 
in  the  functions  call  for,  in  order  to  make  it  commensurate 
with  the  force  expended  in  them.  This  necessity  for  producing 
afflux  of  the  fluids  in  the  organs  simultaneous  with  expan- 
sion, has  forcible  illustration  in  the  lungs,  for  with  the  force 
in  the  chest- walls,  diaphragm,  and  muscles  in  the  abdomen,  in- 
clusive of  the  action  in  the  lungs  themselves,  it  would  be 
utterly  impossible  to  effect  expansion  in  the  absence  of  the 
air ;  which  one  may  readily  prove  upon  himself  by  closing 
the  nose  and  mouth  and  then  attempting  to  inspire.  He 
cannot  expand  his  lungs,  putting  his  force  upon  it  and 
bringing  everything  to  bear.  But  how  easily  it  is  done  with 
removal  of  the  obstruction,  the  air  flowing  into  the  organs 
pari  passu  with  expansion  for  filling  the  room  this  effects  in 
the  alveoli,  pressing  against  the  walls,  and  in  this  manner  aid- 
ing expansion.     So,  likewise,  in  the  capillaries  we  have  the 


17J  .MULTIPLE    DICRoTlSM. 

blood  flowing  into  them  pari  passu  with  expansion  for  occu- 
pying the  room  thus  made,  pressing  against  the  walls,  thereby 
aiding  expansion,  the  principle  being  the  same  precisely  as  ob- 
tains in  the  lungs. 

Multiple  Di erotism. — This  brings  us  to  the  remaining  diffi- 
culty in  the  tissue  tracings,  notably  muliiple  dicrotism.  As 
we  have  seen,  the  systemic  pumping-action  in  the  lungs  is  let 
down  in  stages  to  the  tissue  territory,  wrhich  should  give  us 
but  one  dicrotic  wave  for  all  the  vessels,  inclusive  of  the  heart ; 
whereas,  in  the  capillary  tracings  we  have  sometimes  two,  and 
occasionally  as  many  as  three  or  even  four  present  (Fig.  03), 
and  which  is  undoubtedly  due  to  interference  on  the  part  of 
the  local  functions  and  the  action  in  the  cells.  And  when  so 
many  dicrotic  waves  appear,  we  knowT  there  is  something 
wrong  in  the  tissues,  or  the  cells  are  not  duly  supplied,  and 
the  circumstance  is  announced  in  this  manner.  In  the  case 
before  us,  in  which  the  capillaries  are  choked  with  venous 
blood,  therefore  cannot  act,  dicrotism  disappears  ;  wThile  in 
the  opposite  condition  of  anaemia,  produced  by  an  application 
of  ice  to  the  inner  side  of  the  arm,  or  along  the  course  of  the 
brachial  artery,  for  effecting  contraction  in  the  vessel,  there- 
by to  diminish  the  flow  of  blood  in  the  tissues  below  the  nor- 
mal amount,  accentuates  dicrotism  (Fig.  50),*  at  once  showing 
that  its  origin  is  in  the  capillary  vessels,  and  due  to  the  dimin- 
ished supplies  in  the  cell-brood.  In  other  words,  the  cell- 
brood  is  suffering,  and  this  is  the  only  means  for  increasing 
circulation.  The  action  on  the  artery  irons  out  the  respiratory 
wave,  but  it  does  not  stop  dicrotism,  for  that  is  the  voice  of 
the  cell-brood,  the  vox  populi,  that  will  not  down. 

Explain  the  phenomena  in  the  tissue  tracings  by  the  action 
in  the  heart !  No,  Monsieur!  You  might  as  well  have  attempted 
to  explain  the  force  in  this  vast  republic  by  the  powers  of  the 
Chief  Magistrate,  who  is  only  a  servant  of  the  people,  and  in 
which  every  individual  is  an  independent  integer,  performing 
work  so  as  to  sustain  himself,  if  it  be  not  further  than  to  eat 
and  sleep,  to  rise  up  and  lie  down  again,  and  attend  to  the 
calls  of  nature — all  the  same,  they  are  his  own  acts  and  essen- 

*Ib.,  Fig.  329,  produced  in  the  same  manner  as  the  preceding,  while  the 
fall  in  the  line  of  tracings  indicates  the  amount  of  shrinkage  in  the  hand. 


AE  NESTING    THE  EESPIRATORY   WAVES. 


173 


crT 


p"  2.   i 
§  || 

P  crq 

5-  c 


rs 


&  p. 


V!    b 
"     S 

p 
>3m 


B    * 
CD     

<!  <<i 

E-  <+ 
o 


Ms 


3  g- 


ffl  B 
CD  ~ 
O     O 


&  2 

Ms    CD 


<3     * 


t°    m- 

&9 


174      THE   HEART   NOT   THE   FORCE   IN   THE   CIRCULATION. 

tial  to  him  ;  at  the  same  time,  his  force  is  joined  with  the  body- 
politic  through  coordination  of  the  multitudinous  parts  in  the 
common  federal  centre  for  the  autonomy ;  so  that  the  whole 
performs  as  though  but  a  single  individual  only,  while  separate 
and  independent  action  obtains  all  along  the  line  down  to  the 
individual  integers,  however  insignificant  they  may  be.  So, 
likewise,  in  the  autonomy  in  the  body  itself,  of  which  the  other 
is  reflex  (for  a  stream  cannot  rise  higher  than  its  source),  every 
individual  cell  is  an  integer,  performing  work  relating  to  its 
own  existence  and  the  public  weal,  while  all  the  multitudinous 
parts  are  coordinated  by  means  of  the  correlation  of  the  nerves 
and  nervous  centres  in  the  medulla  oblongata,  the  federal  centre 
of  nervous  force  for  the  organism,  while  separate  and  inde- 
pendent action  throughout  is  a  sine  qua  non  to  existence  :  in 
the  one  not  more  than  in  the  other.  It  follows  that  the  cells 
and  tissues  must  control  their  own  circulation  in  order  to  make 
it  commensurate  with  the  physiological  requirements  and  sus- 
tain existence.  And  had  the  heart  fifty  times  the  force  it  has,  it 
would  not  serve  the  purpose  of  the  circulation,  for  the  principle 
itself  is  wrong.  The  theory  of  Harvey  is  well  enough  under 
limitations,  though  it  was  a  splendid  service  he  rendered  to 
science  in  rounding  the  circulation  through  the  arteries  and 
veins,  sending  it  out  of  the  left  side  through  the  arteries  and 
returning  to  the  right  through  the  veins.  But  then,  Harvey 
knew  nothing  of  the  tissue-circulation,  nor  of  the  vast  cell- 
colonies  upon  its  banks,  and  the  individualism  in  the 
tissues ;  nor  of  the  vast  lymphatic  circulation,  which  is 
slower  than  the  other,  connecting  with  the  nutritive  processes, 
embracing  the  tissue-interstices  and  the  cells,  and  emptying 
into  the  venous  system  at  the  root  of  the  neck ;  nor  of  the 
special  anatomy  in  the  vessels  as  the  relative  anatomical  dis- 
positions for  producing  work  in  the  circulation,  inclusive  of 
the  va  so-motor  system  of  nerves  for  coordinating  them  with 
respiration,  all  of  which  were  subsequent  revelations  effected  at 
the  cost  of  enormous  labors  by  devoted,  self-denying  and 
independent  workers  in  the  field.  Last,  but  not  least,  he 
knew  nothing  of  this  law  of  pressure  underlying  the  organism 
and  forming  the  basis  of  the  entire  mechanics  in  the  body, 
for   Torricelli    had    not    yet  announced  atmospheric    press- 


THE  HKAKT   NOT   THE   FOECE   IX   THE   CTKCULATION.        175 

lire,  coming  out  some  years  after  (1645).  while  Harvey  an- 
nounced his  theory  in  1627,  or  nearly  twenty  years  previously. 
He  was  correct  about  the  blood-circuit,  but  wrong  about  the 
force  being  in  the  heart,  which  cannot  possibly  be  true.  But 
had  it  been  otherwise,  he  could  not  have  worked  out  the 
problem  in  the  absence  of  the  painstaking  anatomical  and 
microscopical  researches,  inclusive  of  the  physiological  experi- 
ments made  upon  the  nerves,  blood-vessels,  etc. ,  which  were 
essential  for  showing  the  relations  they  sustain  to  each 
other  and  their  relevancy  in  the  vital  phenomena  of  which 
respiration  is  the  typical  movement  A  blast  from  John 
Hunter,  ' '  its  heart  is  all  over  its  body,"  *  however,  had  sounded 
the  alarm  all  along  the  line,  and  the  matter  of  the  final  over- 
throw was  only  a  question  of  time,  for  truth  only  can  meet 
and  endure  the  assaults  of  science.  It  is  a  contravention  of  the 
law  underlying  the  organism,  therefore  is  false.  But  we  would 
say  of  these  beautiful  experiments,  that  we  would  not  do 
without  them  if  we  could,  and  we  could  not  do  without  them 
if  we  would 

Circulation  in  the  Tissue-Interstices  and  the  Relations 
which  this  Sustains  to  the  Capillaries  and  Cell-Brood. — For 
illustrating  circulation  in  the  tissue-interstices,  and  for  im- 
pressing it  upon  the  mind,  we  have  prepared  the  following 
illustrative  diagram  of  the  soft  tissues  (Fig.  57).  It  repre- 
sents four  capillary  vessels,  with  the  intervening  tissues  (2), 
formed  of  cells  and  interstices,  sufficient  for  the  purpose  of 
illustration.  Now,  then,  should  the  capillaries  (1,  1,  l,  1) 
contract  for  increasing  pressure  upon  the  blood,  it  would  have 
the  effect  of  increasing  the  distances  between  them,  thereby 
reducing  pressure  in  the  interstices  in  proportion,  and  which 
should  cause  the  liquor  sanguinis  to  escape  through  the  sto- 
mata  into  the  interstices,  which  would  be  from  high  to  low 
pressure.  But  during  expansion  the  opposite  should  take 
place,  since  this  would  have  the  effect  of  diminishing  the 
distances  between  the  capillary  walls,  thereby  increasing  press- 
ure in  the  interstices  in  proportion,  at  the  same  time  devel- 
oping low  pressure  within  themselves  ;  hence,  the  fluids  should 
flow  into  the  vessels  again  from  the  interstices,  being  from 

*  This  remark  was  evoked  by  the  fruitless  search  for  the  heart  of  a  fly. 


176 


CIRCULATION   IN    THE   TISSUE   INTERSTICES. 


high  to  low  pressure,  in  conformity  with  universal  law.  J  n 
other  words,  the  fluids  should  flow  into  and  out  of  the  inter- 
stices from  the  capillaries  during  their  rhythmical  contrac- 
tions and  expansions,  for  the  same  reason  precisely  that  they 
flow  into  and  out  of  the  lungs,  pressure  alike  applying  to  it 
all.  The  blood  corpuscles  being  too  large  to  pass  the  sto- 
mata,  pass  on,  of  course,  into  the  venous  system,  but  yielding 
up  their  oxygen,  which  very  probably  passes  with  the  liquor 
sanguinis  through  the  stomata  to  the  cell-brood,  as  this  would 
greatly  facilitate  the  passage  through  the  membrane.  In  this 
manner,  then,  the  nutritive  and  force-producing  elements  are 
pumped  into  and  out  of  the  interstices  for  the  due  supply 
of  the  cell-brood  ;  and  which  is  readily  increased  or  dimin- 


Fi. 


-An  Ideal  Diagram  of  the  Tissue-Circulation,  showing  the  relations  which  the 
capillaries  sustain  to  the  tissues.     1,  1,  1,  lumen  of  capillary  vessels. 


ished  upon  occasion  in  correspondence  with  the  exigencies  in 
the  functions,  thus  giving  them  complete  control  of  their  own 
supplies,  which  the  scheme  calls  for ;  otherwise  it  must  in- 
evitably fail,  the  sole  purpose  in  all  the  vascular  arrange- 
ments being  the  due  supply  of  the  cell-brood  or  workmen  in 
the  tissues  through  whose  instrumentality  all  the  phenomena 
are  evolved  ;  and  failing  at  one  point,  the  vascular  chain 
would  be  broken,  since  it  all  rests  upon  this  power  of  produc- 
ing rapid  rhythmical  changes  in  pressure  for  increasing  circu- 
lation commensurate  with  the  physiological  requirements, 
no  other  means  applying  for  the  purpose.     Thus  it  is  mani- 


CIECULATIOIS"   I]ST   THE   TISSUE   INTERSTICES.  177 

fest,  for  feeding  the  cell-brood  and  removing  waste  products, 
that  it  calls  for  rhythmical  expansions  and  contractions  in 
the  capillaries,  in  the  arteries,  in  the  heart,  and  in  the  lungs, 
for  producing  an  uninterrupted  now  of  the  fluids  between 
the  cell-brood  and  environment,  whence  all  the  supplies  are 
derived,  and  into  which  the  waste  products  are  returned,  the 
whole  forming  a  connected  movement  in  the  very  nature  of 
things,  since  in  no  other  way  could  a  balance  be  maintained. 
Moreover,  it  is  reduced  to  actual  demonstration  in  the  tissue 
tracings  (Fig.  52,  V)  that  the  tissues  do  expand  and  contract 
regularly  and  rhythmically  synchronous  with  respiration  and 
the  action  in  the  heart,  arteries  and  capillaries,  as  indicated  by 
the  respiratory,  cardo-arterial  and  capillary  waves  which  are 
superposed  upon  each  other  in  the  order  named,  as  before 
remarked,  thus  completing  the  argument  by  accounting  for 
all  the  phenomena  in  the  tissue  tracings  ;  together  with  the 
special  anatomy  in  the  organs,  inclusive  of  the  enormous  web 
of  nerves  for  coordinating  them  with  respiration ;  otherwise 
utterly  inexplicable.  While  all  the  phenomena,  anatomical  and 
physiological,  fall  readily  into  line  in  regular  order  and  succes- 
sion as  appropriate  adjustments  with  the  fundamental  law 
underlying  the  organism,  which  is  thus  being  incessantly  in- 
voked in  the  measure  of  the  requirements,  and  which  is  indi- 
cated by  the  pumping  actions  taking  place  in  the  lungs  and 
organs  of  circulation,  the  whole  rising  and  falling  with  the 
swell  in  the  activities. 

The  following  illustrations  (Pigs.  58,  59)  will  give  some  idea 
of  the  nervous  lines  connecting  the  capillaries  with  the  cell- 
brood,  enabling  them  to  expand  and  contract  the  vessels, 
inclusive  of  the  arterial  feeders  for  increasing  and  diminishing 
the  blood  supply  in  correspondence  with  the  exigencies  in  the 
functions,  aud  which  could  nob  be  effected  in  any  other  way, 
as  they  are  constantly  changing ;  hence,  the  vessels  must  be 
subject  to  the  mandates  in  the  cell-brood.  All  of  which  is 
plain  enough. 

This  force  in  the  cell-brood  for  compelling  supplies  into  the 
body  to  themselves  through  the  action  of  the  rhythmic  centre, 
is  seen  in  the  increase  of  respiration  and  the  action  in  the  heart 
and  arteries  with  the  swell  in  the  activities,  rising  and  falling 


178 


NERVES   CONNECTING   WITH    THE   CELL-BROOD. 


with  these,  the  two  being  in  correspondence  ;  for  in  no  other 
way  could  it  be  effected.  Moreover,  the  influence  which  the 
tissues  exercise  over  circulation  and  respiration  in  abnormal 
conditions,  so  as  to  produce  fever,  notably  traumatism  and 
inflammatory  processes  brought  about  by  any  means,  is  further 
proof  of  the  intimate  connection  subsisting  between  them. 
Finally,  the  accentuation  of  dicrotism  in  the  cardiac  and  arterial 
tracings  by 'anaemia,  superinduced  by  any  cause,  whether  by 
recurrent  hemorrhages  or  exhausting  fevers — e.  </.,  typhoid 
fever,  in  which  the  supplies  are  defective — affords  eloquent 
proof  of  the  power  of  the  cell-brood  for  compelling  the 
rhythmic  centre  to  respond.     One  more  reference,  and  we  shall 


Fig.  58. — Horizontal  Preparation  of  Nictitating  Membrane  of  Frog  in  Chloride  of  Gold, 
showing  the  distribution  of  non-medullated  nerve  fibres  to,  a,  capillary  blood-vessels. 
b.  Coarse  non-medullated  nerve  fibres  giving  off  fine  branches  c,  which  form  a  plexus 
around  the  vessel.     (Oc,  3  ;  obj.,  8.) — Klein. 

have  finished  ;  notably  :  It  is  a  well-known  fact  that  any  cir- 
cumstance which  should  suddenly  reduce  arterial  pressure 
would  accentuate  dicrotism,  bringing  it  out  more  conspicuously 
in  all  of  the  tracings.  Now,  then,  as  arterial  pressure  increases 
circulation  in  the  tissues,  it  follows  that  any  reduction  of 
pressure  would  call  for  corresponding  compensation,  in 
order  to  maintain  circulation  in  the  tissues  up  to  the  norm, 
and  which  can  only  be  done  by  increasing  the  rhythmical 
expansions  and  contractions  in  the  vessels,  and  hurrying  res- 
piration ;  hence  the  quick  respiration  in  these  cases  and  the 


NERVES    CONNECTING   WITH    THE   CELL-BROOD. 


179 


accentuation  of  dicrotism,  which  are  undoubtedly  produced 
through  the  rhythmic  centre  and  reflex  action  propagated  from 
the  cell-brood. 

Concerning  the  Arrest  of  Pulsation  on  the  Distal  Side  of 
Aneurismal  Tumors. — Since  the  central  nervous  system  oper- 
ates the  movements  in  the  vessels,  the  force  being  propagated 

m 


Fig.  59. — Mesentery  of  Frog  prepared  in  Chloride  of  Gold,  showing  the  distribution  of 
non-medullated  nerve  fibres  to  a  capillary  blood-vessel,  a.  b,  A  coarse  non-medul- 
lated  nerve  fibre  giving  off  finer  branches,  which  form  a  plexus  around  the  capillary. 
Some  of  these  finer  fibres  belong  to  the  wall  of  the  vessel.     (Oc,  4  ;  obj.,  8.) — Klein. 

from  the  medulla  oblongata,  or  the  same  as  in  the  voluntary 
movements  with  which  the  vessels  are  coordinated,  this  would 
explain  the  arrest  of  pulsation  in  the  vessels  on  the  distal  side 
of  aneurisms,  which  distends  the  nervous  plexuses,  and  by 


180  CARDIAC   TRACIXGS   DECEPTIVE. 

putting  them  upon  the  stretch  would  inhibit  the  passage  of 
the  currents,  while  the  force  in  the  heart  is  absorbed  in  the 
aneurismal  expansions,  thus  taking  off  nervous  and  cardiac 
force  at  one  and  the  same  time ;  hence,  the  cessation  of  the 
pulsations.  Furthermore,  we  have  seen  that  the  force  in  the 
ventricular  systole  is  needed  for  producing  a  degree  of  press- 
ure in  the  vessels,  in  order  to  enable  them  to  expand,  and  to 
produce  the  energy  involved  in  the  pulse.  So  that  should  the 
nervous  currents  continue  as  before,  the  pulsations  could  not 
be  produced.  Still,  there  is  some  action  in  the  vessels  and  tissues, 
and  circulation  is  sufficient  for  maintaining  life  in  the  affected 
structures,  the  force  in  the  aneurism  itself  not  being  lost,  while 
the  polar  forces  continue  in  action.  In  respect  to  the  fall  in 
arterial  tension,  this,  of  course,  is  an  effort  of  Nature  to  pro- 
mote recovery  by  reducing  pressure  in  the  aneurism,  thereby 
to  lessen  the  tension  so  as  to  diminish  the  danger  of  rupture. 
At  any  rate,  it  has  this  effect,  and  it  cannot  be  accidental,  for 
there  is  method  in  it. 

Concerning  the  Cardiac  Tracings. — The  exceptional  con- 
ditions which  obtain  in  the  heart  tend  to  misdirection  with 
respect  to  the  true  import  of  cardiac  movement  as  registered 
by  the  sphygmograph.  1.  Thus,  being  in  a  manner  suspended 
by  the  vascular  rootlets,  notably  the  pulmonary  and  aortic 
arteries,  resting  lightly  against  the  diaphragm  in  man,  and 
the  sternum  in  quadrupeds,  the  systole  causes  it  to  bound 
against  the  chest  from  the  sudden  action  it  produces  in  the 
vessels,  which  expand  and  elongate,  thereby  throwing  it 
forcibly  against  the  walls,  producing  the  perpendicular  line 
in  the  tracing,  answering  to  diastole  in  the  vessels ;  while 
during  diastole  the  contraction  which  this  produces  in  the 
vessels  pulls  the  organ  from  the  chest  again,  and  so  lets 
down  the  lever  of  the  sphygmograph,  producing  the  oblique 
line  in  the  tracings,  which  answers  to  systole  in  the  vessels. 
Furthermore,  by  reason  of  this  to-and-fro  motion  produced 
in  the  heart,  the  lever  is  incapable  of  following  it  throughout 
the  whole  movement,  the  heart  being  pulled  away  from  it. 
2.  Contraction  in  the  heart  is  from  the  apex  to  the  base,  which 
shortens  and  thickens  the  organ,  in  consequence  pushing  up 
the  lever  of  the  sphygmograph,  thereby  indicating  increase  of 


CARDIAC    TRACINGS   DECEPTIVE.  181 

size  in  the  organ,  whereas  the  organ  is  actually  diminished  in 
size  to  the  extent  of  the  blood  discharged  from  it  during  this 
time ;  hence,  is  deceptive ;  while  during  the  expansion  or 
diastole  the  organ  elongates,  in  this  way  letting  down  the 
lever  again,  thereby  indicating  decrease  of  size,  whereas  the 
size  of  the  heart  is  actually  increased  to  the  extent  of  the 
blood  it  contains  during  this  time ;  hence,  this  is  also  deceptive. 
W]iat  is  desired  in  the  tracings  is  to  get  the  actual  increase 
and  decrease  of  volume  during  diastole  and  systole,  which 
should  give  us  tracings  similar  to  what  obtains  in  the  arteries, 
where  no  such  difficulties  exist,  and  which  the  lever  truly 
registers,  following  the  whole  movement  in  ihe  vessels.  And 
in  order  to  get  them  in  the  heart,  the  instrument  would  have 
to  be  applied  directly  to  the  organ,  as  must  appear  obvious. 
Thus  it  is  seen  the  tracings  are  deceptive.  In  point  of  fact, 
there  are  four  movements  taking  place  in  the  heart,  two  corre- 
sponding to  diastole  and  systole,  and  two  embraced  in  the  to- 
and-fro  movement  corresponding  with  impact  and  recession 
from  the  chest. 

The  Venous  System. — The  arrangements  which  obtain  in  the 
venous  system  are  also  easily  explained.  It  embraces  a  number 
of  circumstances.  In  the  first  place,  it  is  about  four  times  the 
size  of  the  arterial ;  hence,  did  not  some  arrangement  obtain  for 
regulating  the  capacity  by  altering  the  lumen  of  the  vessels, 
the  low  pressure  which  prevails  here,  together  with  the  high 
pressure  in  the  arterial  system,  would  soon  empty  all  the 
"blood  into  the  venous,  bringing  life  to  a  sudden  termination. 
Hence,  the  numbers  of  the  muscles  and  nerves  in  the  veins  for 
effecting  this  circumstance,  while  the  valves  sustain  the  venous 
column  against  gravitation  and  obviate  reflux.  2.  This  in- 
creased capacity  in  the  venous  system  would  explain  the  slow- 
ness of  the  respiratory  rhythm,  and  the  speed  in  the  cardo- 
arterial,  being  as  4  to  1  of  the  former,  the  object  of  respiration 
being  to  bring  the  venous  blood  into  the  alveoli  for  effecting 
oxygenation,  the  heart  and  arteries  serving  to  transfer  it 
thence  to  the  cell-brood  for  evolving  force  in  the  organism, 
while  the  relative  rate  in  the  rhythms  is  necessary  for  main- 
taining a  balance  between  the  venous  and  arterial  systems. 
Hence,  the  nervous  connections  subsisting  between  the  veins 


182  ACTION    IN  THE  VENOUS   SYSTEM. 

and  the  vaso-motor  and  respiratory  centres  for  regulating  the 
venous  system  with  respiration,  for  in  no  other  way  could  a 
balance  be  maintained.  In  line,  respiration,  assisted  by  the 
action  in  the  heart  and  arteries,  pumps  the  blood  out  of 
the  great  venous  reservoir  into  the  alveolar  plexuses  in 
the  measure  of  the  requirements,  while  for  facilitating 
the  action  the  walls  of  the  reservoir  itself  contract  and 
expand  regularly  and  rhythmically,  synchronous  with  res- 
piration. This  would  explain  the  number  of  the  muscles 
in  the  veins,  together  with  the  circular  arrangement  of  the 
muscles  in  the  walls,  or  the  same  as  obtains  in  the  arteries, 
minus  the  thick  elastic  coat  for  producing  the  high  pressure  in 
the  latter,  which  is  not  called  for  in  the  venous  system,  where 
pressure  is  low  in  order  to  give  the  requisite  time  for  effecting 
the  oxygenating  and  nutritive  processes  in  the  tissues,  and  for 
effecting  the  interchanges  in  the  lungs,  which  call  for  a  slow 
circulation,  in  order  to  accomplish  them.  Finally,  the  venous 
system  being  harnessed  to  respiration,  with  the  arterial  for 
producing  an  uninterrupted  flow  of  blood  through  the  circuit, 
it  is  manifest  the  balance  is  struck  in  the  lungs  for  producing 
correspondence  between  the  circulation  of  blood  and  air  in 
the  alveoli,  otherwise  impossible,  in  this  manner,  then,  the 
blood  is  compelled  from  the  centre  to  the  periphery  of  the 
body,  or  from  the  lungs  to  the  tis  ue-territories  through  the 
heart  and  arteries,  thence  back  again  through  the  veins  to  the 
lungs  for  completing  the  circuit,  rocked  in  the  arms  of 
nervous  force,  gently  and  tenderly  nursed  through  the 
work.  Before  leaving  this  portion  of  the  subject,  it  were 
not  amiss  to  refer  to  an  interesting  phenomenon  connected 
with  the  brain  and  venous  system;  notably  the  oscillations 
which  occur  in  the  brain  synchronous  with  respiration,  rising 
up  during  expiration  (as  seen  through  an  opening  in  the  skull), 
and  sinking  down  again  during  inspiration. 

The  explanation  of  this  circumstance  is  furnished  in  the 
alternate  filing  and  emptying  of  the  great  venous  sinuses  at 
the  base  of  the  brain,  and  which  is  due  to  the  mechanics  in 
respiration.  For  example,  during  expiration  there  is  retarda- 
tion of  the  blood  in  the  sinuses,  produced  by  arrest  of  suction- 
force  in  the  chest,  save  in  the  hear,  only,  which  is  not  sufficient 


THE  BRAIN   OSCILLATING  WITH   KESPI RATION.  183 

for  maintaining  an  uninterrupted  flow  of  blood  in  the  sinuses 
— indeed,  nowhere  else  in  the  venous  system,  as  is  fully  proven 
by  the  rapid  accumulations  of  blood  during  prolonged  expi- 
ration, in  which  the  whole  body-surface  becomes  suffused  with 
venous  discoloration  from  venous  stasis  in  the  systemic  capil- 
laries ;  in  consequence,  they  become  distended,  lifting  the 
brain  in  proportion,  causing  it  to  present  at  the  opening 
in  the  skull ;  but  when  inspiration  sets  in,  the  great  increase 
of  suction-force  this  produces  in  the  chest  causes  sudden  efflux 
in  the  sinuses ;  in  consequence,  this  soft  vascular  cushion  col- 
lapses and  as  suddenly  lets  down  the  brain  again. 

In  other  words,  the  matter  resolves  itself  into  one  of  hydraulic 
pressure  simply,  the  brain  riding  up  and  down  upon  this 
hsemal  cushion  underlying  it  with  the  oscillations  of  pressure 
in  the  lungs  and  venous  system.  And  that  it  is  the  correct 
explanation  of  the  phenomenon,  is  proven  by  the  fact  that  the 
motion  does  not  occur  when  the  sinuses  are  laid  open,  in  itself 
furnishing  eloquent  proof  to  the  damming  of  the  blood  in  the 
venous  system  during  expiration,  showing  that  the  heart  is  not 
equal  to  maintaining  circulation  in  the  lungs,  but  a  helpmate 
for  assisting  it  only. 

During  sleep,  therefore,  in  which  respiration  falls  in  fre- 
quency, the  oscillations  are  greatest,  as  a  matter  of  course. 
But  then,  again,  this  subserves  important  functions,  since  the 
plethora  of  the  capillaries  and  the  slowing  of  circulation  which 
takes  place  in  sleep  conduces  to  rapid  nutrition,  since  it  favors 
crystallization,  which  requires  a  slow  circulation.  But  it  is  all 
in  correspondence ;  thus,  with  diminished  respiration  there  is 
diminished  action  in  the  heart  and  arteries,  in  correspondence 
with  this  circumstance,  for  maintaining  a  balance  in  the  arterial 
and  venous  systems,  with  a  slow  circulation  for  promoting  the 
nutritive  processes ;  while  with  revival  of  consciousness  and 
the  activities,  a  simultaneous  increase  occurs  in  correspond- 
ence with  this  circumstance  for  evolving  the  force  expended 
in  the  organism,  extending  through  the  entire  mechanics  in 
wondrous  order  and  harmony,  considering  the  complexity 
which  exists,  which,  of  course,  is  effected  by  means  of  nervous 
force  propagated  from  the  medulla  oblongata,  the  solar  centre 
of  the  organism.    But  when  considered  in  the  light  of  this  com- 


184  CONCLUSION   OF   SYSTEMIC   CIRCULATION. 

mon  law  underlying  it,  it  is  readily  perceived  why  there  should 
be  such  precision  "and  harmony  pervading  it,  and  why  similar 
arrangements  should  obtain  in  all  the  animals,  commensurate 
with  the  stage  in  development ;  and  when  one  reflects  over 
the  circumstance  that  everything  is  regulated  by  law.  it  is 
passing  strange  that  this  fact  had  not  been  applied  to  the 
animal  circulation. 

Turn  we  now  to  the  portal  circulation  and  the  mechanics  in 
the  abdomen  for  further  corroborative  evidence  of  this  funda- 
mental law  underlying  the  organism,  with  which  everything 
must  have  adjustment,  as  has  already  been  remarked. 


CHAPTER  IX. 

THE    POETAL    CIRCULATION"  AND  THE    MECHANICS    IN"    THE 

ABDOMEN. 

The  Intestines  of  Mammalia  Filled  and  Distended  with  Air  for  Increasing  the  Digestive 
and  Absorptive  Processes,  Serving  the  Purpose  of  an  Elastic  Cushion  for  Trans- 
mitting the  Force  in  the  Gut  upon  the  Food  for  the  Purpose— The  Action  not  Un- 
like that  which  Obtains  in  a  Churn— Necessity  for  a  Diaphragm— Existence  of  High 
Pressure  in  the  General  Cavity  of  the  Abdomen  Produced  by  the  Air  in  the  Intes- 
tines for  Increasing  the  Portal  Circulation  so  as  to  Maintain  this  in  Correspondence 
with  the  Absorptive  Processes— Principle  of  Coordination  Applied  to  the  Stomach 
and  Walls  of  the  Abdomen  for  Effecting  Ingestion  and  Office  of  the  Pneumogastric 
and  Phrenic  Nerves  in  Connection  Therewith— Why  the  Animal  Rises  when  Eating 
and  Drinking— Why  Respiration  is  Suspended  During  Deglutition  or  After  the  Ali- 
ment has  Passed  the  Glottis,  and  the  Danser  of  Intrusion  in  the  Air-Passages  is  Over— 
The  Action  in  the  Stomach  and  the  Physiological  Anatomy  in  the  Organ— The  Role 
in  the  Air-Cus  ion  in  Connection  Therewith— Similar  Survey  of  the  Intestines— 
The  Rapid  Absorption  of  Fat,  Alcohol  and  Other  Non-Dialyzable  Substances  Easily 
Explained— The  Fine  Adjustments  in  the  Museularis  Mucosoz  and  the  Mode  of 
Coordinating  Them  with  the  Muscles  in  the  Walls  for  Making  Absorption  Very 
Effective— No  Difficulty  in  Absorption. 

Having  passed  in  rapid  review  tile  phenomena,  anatomical 
and  physiological,  appertaining  to  the  systemic  circulation, 
and  their  relevancy  to  the  law  underlying  the  organism,  fully 
shown  and  established  by  indisputable  evidence,  we  are  now 
prepared  to  take  up  the  portal  circulation  and  the  mechanics 
in  the  abdomen  for  compelling  the  digestive  and  absorptive 
processes  to  be  commensurate  with  the  physiological  require- 
ments, so  as  to  maintain  a  balance  in  the  organism  ;  otherwise 
impossible.  The  adjustments  which  obtain  in  this  respect  are 
beautiful  to  look  upon,  and  passing  all  belief,  could  they 
not  be  inspected  with  one's  own  eyes.  But  there  they  are, 
true  enough,  and  open  to  inspection  And  taken  all  in  all, 
the  number  and  diversity  of  the  adjustments,  the  charming 
manner  of  the  workmanship,  and  the  mechanics  in  the  ab- 
domen exceed  anything  in  the  domain.  The  finest  adjustments 
come  in  with  the  nervous   apparatus  for  effecting  the  mul- 


186  OFFICE    OF  ATIt  IN    THE  INTESTINES. 

titudinous  actions  in  the  organs,  and  for  coordinating  them 
with  respiration,  the  whole  being  intimately  interwoven  with 
the  systemic  mechanics  and  blending  thoroughly  with  this. 
We  shall  not  dwell  longer  than  necessary,  passing  over  the 
structures  rapidly. 

Antecedent  to  the  mammalia,  the  intestines  remain  small 
and  contracted,  the  walls  comparatively  thick,  and  the 
cavity  filled  with  liquid  contents  (Fig.  60) ;  but  with  the 
mammalian  stage  in  development  the  organs  are  widely  ex- 
panded, the  walls  comparatively  thin,  while  the  cavity  is 
filled  and  distended  with  air,  the  liquid  contents  at  the  bot- 
tom and  around  ^he  sides  of  the  tube  (Fig.  61,  1,  2).  Acci- 
dental !  No  ;  of  course  not !  Nature  does  not  deal  in  acci- 
dents ;  her  arrangements  are  all  methodical,  being  based  upon 
law,  here  as  elsewhere. 

Besides,  accidents  would  not  be  confined  to  one  class  of 
animals  only,  all  the  rest  escaping.  And  if  you  mean  that 
the  air  in  the  intestines  of  mammalia  is  the  result  of  fermenta- 
tive changes  in  the  food,  or  to  decomposition,  you  are  entirely 
wrong ;  the  animals,  moreover,  live  on  the  same  kind  of 
food  as  all  the  others,  namely,  vegetable  and  animal  food — 
some  one  kind  only,  others  both  kinds  ;  so  that  in  no  sense 
can  there  be  any  excuse  for  such  opinion,  and  it  is  totally 
unscientific. 

Briefly,  the  explanation  for  this  phenomenon  is  as  follows : 

1.  It  affects  a  great  increase  in  the  mucous  surface,  thereby 
increasing  the  digestive  and  absorptive  processes  correspond- 
ingly, and  which  is  necessary  for  evolving  the  force  which  is 
expended  in  them  and  for  maintaining  a  balance  in  the  organ- 
ism otherwise  impossible. 

2.  By  means  of  this  air  in  the  intestines,  the  force  in  the 
walls  of  the  organs  is  applied  to  the  aliment  for  compelling 
absorption  to  be  commensurate  with  the  physiological  re 
quirements ;  at  the  same  time  it  increases  the  digestive 
processes  correspondingly,  by  the  free  movement  it  effects 
in  the  food,  bringing  it  into  rapid  contact  with  the  secre- 
tions that  are  poured  out  upon  the  mucous  surface,  and 
which  is  produced  by  the  action  of  the  nerves  and  muscles 
in  the  walls  of  the  organs  from  sensory  impressions  in  the 


OFFICE    OF    AIR    IN    THE   INTESTINES. 


187 


mucous  membrane  produced  by  the  food.  In  short,  the  me- 
chanical principle  is  the  same  as  obtains  in  the  case  of  residual 
air  in  the  lungs,  for  compelling  circulation  of  the  blood  to  be 
in  correspondence  with  the  circulation  of  air  in  the  alveoli  ; 
only  the  air-chamber  does  not  leak,  as  in  the  former  case,  the 
air  being  all  retained,  while  the  force  in  the  gut  is  available 
for  compressing  the  elastic  cushion  against  the  aliment  for 

A 

JP 

— fflfc- 

.3 

Fig.  60.— A  Diagrammatic  Transverse  Section  of  the  Intestines  in  Aves,  Reptilia,  Pisces, 
etc. ,  showing  the  relative  difference  in  the  size  of  the  lumen  with  Mammalia,  together 
with  the  circumstance  that  the-  organs  are  filled,  with  liquid  contents,  simply. 

B 


Fig.  61. — A  Diagrammatic  Transverse  Section  of  the  Intestines  in  Mammalia,  showing 
the  great  increase  in  the  lumen  of  the  organ  and  the  large  amount  of  air  it  contains. 
1,  liquid  contents  at  the  bottom  and  diffused  around  the  sides  ;  2,  air-cushion  ;  3, 
peritoneum,  longitudinal  muscles  adjacent ;  4,  circular  muscles  ;  5,  mucous  mem- 
brane. 

compelling  absorption  and  digestion  to  be  in  correspondence 
with  the  physiological  requirements  ;  otherwise  impossible. 
Moreover,  it  affords  a  ready  explanation  of  the  special  anatomy 
in  the  organs  as  means  to  ends,  otherwise  inexplicable  ;  nota- 
bly, the  arrangements  in  the  muscles  and  nerves  for  effecting 
the  changes  of  pressure  in  the  contents  in  the  gut,  which  we 


188  OFFICE    OF   AIR    IN   THE  INTESTINES. 

will  come  to  further  on.  Finally,  this  would  account  for  the 
rapid  evolution  of  gas  in  morbid  conditions  of  the  organs  ;  e.  g. , 
dyspepsia,  colic,  hysteria,  in  the  entire  absence  of  food-changes, 
such  as  are  involved  in  decomposition  or  fermentation,  the 
food  acting  only  as  an  irritant,  and  thereby  disturbing  the 
balance  in  the  secretory  processes  in  the  organs,  and  thus 
causing  the  evolution  of  gas  in  abnormal  proportions. 

:i.  Last,  but  not  least,  by  means  of  the  air  in  the  intestines, 
the  increase  of  pressure  is  produced  within  the  general  cavity 
of  the  abdomen  for  producing  correspondence  between  the  por- 
tal circulation  and  the  absorptive  processes  in  the  intestines, 
the  force  in  the  walls  of  the  abdomen  being  available  for  the 
purpose  of  increasing  the  pressure  which  occurs  during  res- 
piration, especially  during  inspiration,  when  the  diaphragm 
descends  ;  a  circumstance  which  it  would  be  difficult  to  over- 
estimate, being  necessary  for  maintaining  a  balance  in  the  circu- 
lation. But,  before  proceeding  farther,  it  were  well  to  demon- 
strate this  circumstance,  that  it  may  rest  upon  something 
better  than  mere  assertion.  For  this  purpose,  a  dog  was 
chosen,  and,  placing  it  under  chloroform,  I  made  use  of  an 
abdominal  sound  (Fig.  62),*  improvised  for  the  purpose,  by 
means  of  which  1  ascertained  the  following  facts  : 

1.  The  existence  of  high  pressure  in  the  general  cavity  of 
the  abdomen,  produced  by  the  gas  in  the  intestines.  2.  That 
pressure  is  increased  during  inspiration,  leaping  up  the  in- 
stant it  sets  in,  and  continuing  higher  than  the  norm  through 
the  whole  period.  That  pressure  is  lower  during  expiration 
than  inspiration,  expansion  in  the  diaphragm  being  so  very 
rapid  as  to  prevent  any  diminution  in  the  abdominal  area  by 
the  contraction  of  the  muscles  in  the  abdomen,  otherwise  in- 
evitable, and  there  can  be  no  doubt  that  it  is  a  normal  physio- 

*  The  instrument  was  made  as  follows  :  The  end  of  a  glass  pipette  was 
passed  into  a  small  metallic  bulb  perforated  with  holes,  over  which  was  drawn 
a  gum-elastic  bag  and  firmly  secured  by  means  of  turns  of  waxed  thread.  The 
instrument  is  then  filled  with  water  till  it  mounts  half-way  up  the  6tem 
and  tightly  corked  to  prevent  escape  when  being  passed  into  the  cavity  of  the 
abdomen. 

When  the  water  is  poured  into  the  open  end  of  the  stem  it  distends  the  gum 
bag,  while  the  bulb  prevents  it  all  from  being  driven  out  under  pressure.  It  is 
a  rude  device,  but  answers  the  purpose  very  well. 


HIGH   PRESSURE   EN"   THE  ABDOMEN. 


189 


logical  condition  to  meet  the  special  requirements  in  the  portal 
circulation  ;  the  obvious  purpose  being  to  obviate  strain  to> 
the  heart  and  vense  cavse  by  reason  of  the  damming  of  the 
blood  in  the  right  side  during  expiration,  as  has  already 


Fig.  62. — Abdominal  Sound. 

been  demonstrated.  I  had  no  means  of  ascertaining  it  in 
millimeters,  but  the  intra-abdominal  pressure  is  very  con- 
siderable. Thus,  when  the  sound  was  first  introduced  through 
an  opening  in  the  linea  alba,  immediately  above  the  umbilicus,, 


190  HIGH   PRESSURE   IN   THE  ABDOMEN. 

and  the  cork  withdrawn  from  the  stem  so  as  to  permit  the 
water  to  escape,  it  spurted  out  with  great  force  against  the 
opposite  wall  some  eight  feet  distant,  and  would  have  gone 
farther,  striking  it  some  feet  from  the  ground. 

It  rose  and  fell  with  inspiration  and  expiration,  and  had 
there  been  sufficient  water  in  the  instrument,  would  have  issued 
per  saltam  with  respiration,  the  same  as  arterial  blood  with 
the  rhythms  in  the  heart  and  arteries,  showing  the  great  press- 
ure in  the  abdomen  and  the  inevitable  effect  it  must  have  upon 
the  venous  system  in  the  abdomen.     I  do  not  mean  that  the 
pressure  is  as  great  as  in  the  arterial  system,  but  that  there  is 
considerable  pressure  in  the  abdomen,  which  oscillates  with 
respiration,  but  highest  during  inspiration.    After  the  first  jet, 
other  smaller  jets  followed  synchronous  with  inspiration,  till  at 
last  the  quantity  of  water  was  so  reduced  that  it  simply  rose  and 
fell  in  the  instrument,  shooting  up  the  stem  during  inspiration, 
then  falling  back  out  of  sight  again  during  expiration  into 
the  bulb  of  the  instrument,  the  delicate  india-rubber  bag  allow- 
ing it  to  do  so.     And  it  thus  continued  to  oscillate  all  the  while, 
no  matter  where  the  bulb  was  shifted,  up  or  down,  in  the  pel- 
vis as  well  as  elsewhere,  but  always  greatest  in  the  region  of  the 
diaphragm  and  stomach.     Furthermore,  the  pressure  is  higher 
in  2,  fat  than  in  a  lean  dog,  showing  conclusively  that  there  is 
greater  difficulty  in  carrying  on  the  portal  circulation  in  the 
former  condition,  since  the  accumulation  of  fat  in  the  viscera 
tends  to  embarrass  respiration,  thereby  choking  the  portal 
vessels ;   hence  this  circumstance.      Fat  animals  are  always 
windy,  the  gas  frequently  escaping  by  the  anus  from  the  press- 
ure in  the  abdomen      The  explanation  of   the  mechanics  is 
obvious  enough.     The  gas  acts  as  a  lever  under  the  force  of 
the  muscles  in  the  abdomen,  and  in  the  walls  of  the  intestines 
for  overcoming  inertia  in  the  venous  blood,  in  this  manner  lifting 
it  up  to  the  chest-cavity  and  lungs,  inclusive,  of  course,  of  the 
liquid  aliment  with  the  portal  blood,  the  whole  forming  a  con- 
nected movement,  and  it  must  do  so  in  the  very  nature  of  things. 
It  is  all  very  beautiful,  while  nothing  could  be  more  simple  : 
air  upon  the  one  hand,  muscular  force  upon  the  other,  the 
blood  with  the  nutritive  and  force-producing  elements  in  the 
intestines  sandwiched  between  them,  with  the  nervous  force 


HIGH   PKESSUEE  IN   THE  ABDOMEN.  191 

in  the  medulla  oblongata  for  operating  the  mechanics  and  co- 
ordinating it  with  respiration.     Mlrdbile! 

The  pressure  in  the  abdomen  would  account  for  the  eleva- 
tion of  the  diaphragm  in  the  chest-excavation,  as  has  already 
been  shown  (Fig.  27).  It  is  higher  after  death  by  reason  of 
the  contraction  in  the  lungs,  forcing  out  a  larger  amount  01 
residual  air  than  during  life,  while  the  viscera  in  the  abdomen 
are  compelled  into  the  excavation  in  order  to  equalize  press- 
ure ;  hence  this  circumstance. 

And  it  would  also  account  for  the  tendency  in  the  viscera  to 
escape  through  the  natural  and  artificial  openings  from 
wounds  in  the  abdomen,  and  which  is  increased  during  in- 
spiration,* loops  of  intestine  and  portions  of  omentum  fairly 
leaping  through  the  openings  the  instant  they  are  made,  from 
the  high  pressure  in  the  abdomen.  It  takes  considerable  force 
to  cause  the  viscera  to  burst  through  the  openings  in  this 
manner.  Last,  but  not  least,  when  the  diaphragm  is  rup- 
tured or  incised  (as  has  occasionally  occurred),  the  viscera  rush 
through  the  opening  into  the  pleura  and,  compressing  the 
lungs,  extinguish  life.  In  conclusion:  That  there  is  high 
pressure  within  the  intestinal  cavity  itself  also  admits  of  easy 
demonstration  by  simply  puncturing  the  gut,  when  the  gases 
escape  with  audible  noise,  the  walls  at  the  same  time  col- 
lapsing, showing  conclusively  that  they  had  been  distended 
by  the  gases.  Moreover,  it  would  explain  the  rotund  appear- 
ance, not  to  be  accounted  for  by  any  other  hypothesis,  the 
walls  being  thus  ballooned  by  the  air. 

We  are  now  prepared  to  take  up  the  mechanics  in  the  in- 
testinal canal.  As  we  live  by  breathing  and  eating,  the  first 
thing  to  note  is  the  adjustment  that  obtains  between  eating 
and  breathing,  an  important  adjustment  obtaining  in  this 
respect.  Thus,  when  the  animal  desires  to  take  food  (solids  or 
liquids,  it  matters  not),  it  rises  to  its  feet  (man  to  the  sitting 
posture)  in  order  to  do  so.  Second,  that  during  deglutition., 
respiration  is  suspended.  The  explanation  of  the  phenomena 
is  sufficiently  easy. 

*  A  very  notable  example  in  the  experience  of  obstetricians  is  the  gush  of 
blood  which  precedes  the  outcry  of  the  parturient  woman,  and  which  is  pro- 
duced by  inspiration. 


192  SUSPENSION   OF   ELSPIUAIION   IN   DEGLUTITION. 

It  rises  up  in  order  to  reduce  pressure  and  facilitate  expan- 
sion in  the  abdomen,  which  are  essential  for  introducing  the 
food  into  the  cavity,  doing  so  instinctively,  but  first  com- 
pelled to  it.  At  the  same  time,  gravitation  is  also  brought  to 
bear  for  facilitating  expansion,  acting  also  upon  the  food ; 
whereas  in  the  recumbent  posture  it  acts  adversely  by  increas- 
ing pressure  in  the  abdomen,  from  the  weight  in  the  body  com- 
pressing the  abdomen  against  the  ground,  thereby  inhibiting 
expansion  ;  a  circumstance  which  is  also  easily  demonstrated 
by  lying  upon  the  abdomen,  then  making  an  attempt  to  swallow 
liquid  or  solid  food.  It  cannot  be  done  but  to  the  most  limited 
extent ;  and  if  the  abdomen  be  at  all  pendulous,  it  will  be 
utterly  impossible  to  get  down  any  whatever.  This,  then,  is 
the  explanation  for  the  rising  up  to  eat  and  drink. 

2d.  The  suspension  of  respiration  has  its  explanation  in  the 
circumstance  that  the  diaphragm  must  be  relaxed  (expanded) 
in  order  to  reduce  pressure  in  the  abdomen  and  relieve  the 
muscular  constriction  around  the  oesophageal  opening  in  the 
diaphragm,  which  constricts  the  tube  during  inspiration. 
Hence,  there  must  be  no  inspiratiory  effort  during  deglutition. 

It  is  the  common  impression,  however,  that  ''suspension  of 
respiration  during  deglutition  is  designed  to  obviate  intrusion 
of  the  food  into  the  air-passages  ;' '  but  while  this  important 
end  is  also  attained,  it  is  not  the  controlling  principle  in  the 
mechanics,  as  is  fully  proven  by  the  fact  of  its  continuance  after 
the  food  has  passed  the  glottis  and  when  all  danger  from 
intrusion  is  over,  not  pausing  till  the  bolus  has  traversed  the 
entire  distance  in  the  gullet  and  been  safely  lodged  within  the 
stomachal  cavity.  And  the  glottis,  being  at  the  very  com- 
mencement of  the  gullet,  is  quickly  passed,  from  the  energetic 
action  of  the  muscles  of  the  pharynx,  which  are  of  the  striated 
variety  for  compelling  rapid  action,  the  evident  purpose  being 
to  rush  it  by  the  glottis,  while  it  is  the  distance,  together  with 
the  slower  movements  in  the  gullet,  that  consumes  the  time. 
Nor  is  the  temporary  arrest  of  the  bolus  in  the  gullet  from 
mechanical  causes,  with  resumption  of  respiration  before  the 
act  is  completed,  an  infraction  of  this  important  principle, 
since  for  the  final  consummation  it  still  requires  the  suspension 
of  respiration,  and  as  though  there  had  been  no  stoppage 


THE  ACTION    IN   DEGLUTITION.  193 

whatever,  in  order  to  relax  the  oesophageal  opening,  or  pillars, 
in  the  diaphragm,  at  the  same  time  reducing  intra-abdominal 
pressure,  the  whole  diaphragm,  together  with  the  muscles 
in  the  abdomen,  expanding,  which  is  accomplished  by  means 
of  the  reflex  actions  set  up  through  the  pneumogastric  nerves 
from  sensory  impressions  in  the  mucous  surface. 

Thus,  deglutition  is  seen  to  be  far-reaching,  a  movement,  in 
short,  extending  from  centre  to  circumference  of  the  body,  and 
involving  the  most  extensive  adjustments  for  effecting  it,  while 
the  mechanical  principle  underlying  it  is  the  one  of  pressure, 
and  the  power  of  producing  rapid  rhythmical  changes  in 
pressure.  In  the  quadruped,  when  drinking  from  the  ground, 
the  liquid  has  to  ascend  the  tube  ;  hence,  it  involves  a  greater 
expenditure  of  force  on  the  part  of  the  gullet  than  in  man, 
who  lifts  it  to  his  lips,  thereby  gains  the  force  in  gravitation. 
But  the  juggler,  drinking  while  standing  upon  his  head,  re- 
verts to  the  old  mechanics,  and  while  he  is  able  to  accomplish 
the  feat,  he,  at  the  same  time,  finds  it  much  more  difficult  than 
the  natural  way,  or  by  lifting  it  to  his  lips  with  a  cup,  with  the 
body  in  an  upright  position,  or  with  the  head  up.  But  it  is 
quite  as  easy  to  effect  expansion  in  the  stomach  and  abdomen 
as  in  the  normal  position.  The  popular  interest  it  excites  is 
amusing.  The  idiots !  The  act  of  deglutition,  however,  is 
somewhat  complex.  Briefly,  when  the  food  has  been  duly  in- 
salivated and  all  is  ready,  the  mouth-cavity  contracts  upon  its 
contents,  the  tongue  being  forced  up  against  the  hard  palate 
from  before  backward  by  the  action  of  the  hyo-glossal,  stylo- 
glossal,  and  palato-glossal  muscles,  strongly  arching  the  organ 
upon  itself  so  as  to  throw  the  convex  surface  against  the  sur- 
face of  the  hard  palate,  and  pulling  it  from  before  backward ; 
at  the  same  time,  the  larynx  is  firmly  approximated  against 
the  base  of  the  tongue  by  the  action  of  the  genio-hyoidei, 
thyro-hyoidei,  mylo-hyoidei,  and  the  anterior  bellies  of  the 
digastric  muscles,  the  effect  being  to  close  the  glottis  and 
effectually  prevent  intrusion  of  the  food  into  the  air-passages, 
the  base  of  the  tongue  also  arching  over  it.  At  the  same  time 
this  is  going  on,  however,  changes  are  taking  place  in  the  back 
of  the  mouth  and  pharynx  for  accelerating  the  passage  of  the 
bolus,  notably  as  the  root  of  the  tongue  approaches  the  velum 


194  THE   ACTION    IN   DEGLUTIT  ON. 

palati  and  fauces — previously  tightly  closed ;  these  expand, 
the  velum  being  drawn  upward  and  backward,  and  op- 
posed to  the  posterior  wall  of  the  pharynx  by  means  of 
the  pharyngo-palati,  and  the  levator  and  circumflexus  pal- 
ati for  obviating  intrusion  in  the  nasal  passages,  and  with 
the  pharynx  at  the  same  time  widely  expanding  for 
sucking  the  food.  The  elevation  of  the  larynx  must  have 
this  effect,  not  to  mention  the  special  action  in  the  mus- 
cles ;  the  bolus  readily  passes  into  the  excavation,  which 
closes  at  once  upon  it ;  thence,  by  a  series  of  rhythmical  ex- 
pansions and  contractions,  is  compelled  through  the  tube  to 
the  stomach.  And  the  pharyngeal  muscles  being  striated, 
this  secures  energetic  action  in  the  pharynx  for  rushing  it  out 
of  the  dangerous  locality,  as  before  remarked.  Otherwise, 
this  differentiation  of  the  muscles  in  the  gullet  is  inexplicable. 
As  soon  as  the  food  reaches  the  pharynx,  the  rellex  actions 
are  at  once  set  up,  nor  pause  till  the  bolus  has  reached  the 
stomach.  The  sensory  fibres,  which  lead  to  the  rellex  move- 
ments in  the  gullet,  are  contained  in  the  palative  branches  of 
the  fifth,  and  in  the  pharyngeal  branch  of  the  pneumogastrics, 
and  the  centre  for  their  movements  is  in  the  olivary  bodies  in 
the  medulla  oblongata  (Schroder  v.  d.  Kolk).  That  the  pro- 
gressive contractions  and  expansions  which  occur  in  the 
oesophagus  during  deglutition  do  not  depend  upon  the  stimu- 
lation of  the  advancing  bolus,  but  are  the  result  of  a  central 
coordination,  is  evident  from  the  fact  that  the  wave  travels  over 
ligatures  or  even  excised  portions  of  the  oesophagus  (Mosso). 
One  other  thing  in  this  connection  :  When  the  mouth  is 
closed  it  has  a  negative  pressure  corresponding  to  2  to  4  milli- 
meters of  mercury  (Mezzer).  In  this  manner,  then,  atmos- 
pheric pressure  would  serve  for  supporting  the  maxilla  and 
tongue,  while  for  increasing  it,  for  producing  the  suction-action 
in  the  cavity  during  nursing  and  drinking,  the  lips  close  around 
the  mamma  or  open  in  the  fluid  ;  at  the  same  time,  the  cavity 
is  expanded  by  expanding  the  buccinator  muscles  and  depress- 
ing the  tongue,  the  fluids  flowing  in  till  expansion  ceases, 
when  the  cavity  contracts  firmly  upon  it  for  compelling  it  into 
the  pharynx,  etc.,  etc.  First  and  foremost,  however,  we  must 
keep  conspicuously  in  the  foreground  the  principle  of   co- 


STOMACH  AND  WALLS  OF  THE  ABDOMEN  COOEDINATED.    195 

ordination  as  it  relates  to  the  internal  and  external  parts, 
together  with  the  nervons  combinations  in  the  mednlla 
oblongata  for  compelling  simultaneous  action  in  them,  since 
this  is  essential  to  the  mechanics  and  the  performance  of  the 
functions  in  the  abdomen. 

As  before  remarked  (p.  60,  Fig.  16,  2,  3),  the  waves  of  ex- 
pansion in  deglutition  are  not  lost,  but  are  added  up  in  the 
general  expansion  which  takes  places  in  the  stomach  and  the 
cavity  of  the  abdomen,  all  the  parts  expanding  pari  passu 
with  the  ingestion  of  food  ;  otherwise,  it  were  utterly  impos- 
sible to  introduce  the  food,  since  the  progressive  increase  in 
pressure  this  should  occasion  in  the  stomach  and  abdomen 
would  inevitably  produce  regurgitation.  The  extension  of 
the  pneumogastric  trunks  to  the  stomach,  which  they  literally 
envelop  with  nervous  filaments,  and  the  correlation  of  these 
nerves  with  the  nerves  to  the  muscles  in  the  abdomen,  nota- 
bly the  intercostals  and  phrenic  nerves,  has  its  explanation  in 
this  circumstance ;  for  in  no  other  way  could  correspondence 
be  produced  for  maintaining  a  balance  in  pressure  in  the  stom- 
ach and  abdomen,  since  the  stomach  is  separated  from  the 
walls  by  the  differentiation  of  a  peritoneal  cavity. 

Furthermore,  that  the  abdomen  does  actually  expand  pari 
passu  with  ingestion  is  matter  of  easy  demonstration  by 
taking  the  dimensions  before  and  after  a  meal,  when  it  may 
be  at  once  seen  that  expausion  is  in  correspondence  with  the 
amount  of  ingesta.  That  ends  the  matter  there.  ISTow,  then, 
it  will  scarcely  be  contended  that  deglutition — that  is,  the 
action  in  the  gullet — can  force  open  the  stomach  in  order  to 
produce  the  increase  in  the  size  of  the  organ  when  thus  dis- 
tended with  food,  the  powerful  muscles  in  the  abdomen  in- 
hibiting this  circumstance,  not  to  mention  the  strain  it  would 
involve  to  the  stomach.  It  sends  a  spasm  through  the  chest. 
Gott  in  Himmel !  Give  it  the  go-by  instantly ;  it  would  choke 
the  life  out  of  you.  And  do  not  think  of  forcing  open  the 
stomach  and  abdomen  by  any  means  whatever,  but  rather 
permit  them  to  expand  of  their  own  option  under  the  stimulus 
of  the  food  and  the  reflex  actions  this  excites  in  the  medulla 
oblongata  from  the  sensory  impressions  it  produces  in  the 
mucous  surface,  or  the  same  as  obtains  in  the  lower  animals. 


196 


PHYSIOLOGICAL  ANATOMY. 


You  cannot  improve  the  mechanics,  founded  as  it  is  in  the 
organic  laws,  therefore  wonderfully  perfect  and  harmonious 
in  all  the  movements. 

In  short,  the  effort  to  explain  this  beautiful  mechanism  in 
the  absence  of  the  fundamental  principle  underlying  it,  is  as 
futile  as  the  attempt  to  sit  upon  nothing.  Food  requires 
room  to  be  made  for  it,  and  you  must  permit  the  stomach  and 
abdomen  to  expand  pari  passu  with  the  ingoing  of  the  ingesta 
in  order  to  make  this  room ;  otherwise,  the  food  could  never 
be  introduced  and  animal  life  would  be  impossible.     All  is 


Tig.  63.— A  Longitudinal  Section  of  the  Human  Stomach,  showing  the  folds  in  the 
mucous  membrane. — Gray. 

chaos  in  the  absence  of  this  law  and  what  it  involves.     It  is 
needless  to  extend  the  matter. 

Concerning  the  Action  in  the  Stomach. — Coming  to  the  stom- 
ach, the  first  thing  to  engage  attention  is  the  anatomical  dis- 
positions in  the  mucous  membrane,  which  contains  the 
organs  of  secretion  and  absorption.  In  the  empty  or  con- 
tracted condition  of  the  stomach,  the  mucuous  membrane  is 
thrown  into  a  series  of  longitudinal  folds  (Fig.  63),  the  loose 
areolar  tissue  connecting  it  with  the  adjacent  circular  mus- 


PHYSIOLOGICAL    ANATOMY. 


197 


■cles,  which  form  a  continuous  cylinder  through  the  whole 
length  of  the  intestinal  canal,  admitting  of  this  action  in  the 
mucous  membrane,  or,  rather,  compelling  it,  since  it  does  not 
possess  the  same  powers  in  expansion  and  contraction  as  the 
muscles  ;  hence,  when  the  organ  becomes  greatly  contracted, 
as  in  the  empty  condition,  this  folding  in  the  mucous  lining 
occurs,  while  in  the  expanded  condition,  produced  by  inges- 


Fig.  64. — A  Transverse  Section  through  the  Fundasof  the  Stomach  in  a  Child. — Verson. 
a,  a,  cylindrical  epithelium  ;  b,  b,  peptic  tubes  ;  c,  muscalaris  mucosas  ;  d,  sub- 
mucous tissue  ;  e,  circular  muscular  layer  ;  g,  peritoneum  ;  h,  ganglia  of  Auerbach. 

tion,  they  again  apply  themselves  accurately  to  the  muscles, 
causing  the  gland-tubes  to  stand  erect  and  facilitating  circula- 
tion and  secretion  correspondingly.  The  following  beautiful 
cut,  by  a  distinguished  German  microscopist,  will  serve  for 
impressing  the  matter  (Fig.  64).  The  great  relative  thickness 
of  the   circular  muscles   (e)  is   a   conspicuous   circumstance, 


198  PHYSIOLOGICAL    ANATOMY. 

while  the  loose  areolar  tissue  is  drawn  into  forcible  extension 
by  the  folds  in  the  mucous  membrane.  Of  course,  the  secre- 
tions for  dissolving  the  food  should  be  compelled  out  of  the 
tubes  as  fast  as  they  are  formed  ;  and  looking  to  the  secretory 
processes  in  these  organs,  then,  it  is  readily  perceived  how  this 
is  materially  assisted  by  the  action  in  the  circular  muscles  for 
compelling  the  contents  in  the  cavity  ;  at  the  same  time,  addi- 
tional force  is  put  upon  it  by  means  of  the  finer  adjustments 
that  obtain  in  the  muscalaris  mucosa. 

Briefly,  they  are  as  follows  :  The  peptic  and  mucous  glands 
occupy  a  perpendicular  position  in  the  mucosa,  the  closed 
end  resting  against  the  circular  muscles,  with  the  loose  con- 
nective tissue  and  muscalaris  mucosae  between,  in  intimate 
relation  with  them,  the  open  end  at  the  free  surface.  The 
result  of  this  arrangement  is  that,  when  the  stomach  contracts 
upon  its  contents  (inclusive  of  the  air  it  contains  for  in- 
creasing the  action)  it  compresses  them  firmly  against  the 
mucous  surface,  tending  to  flatten  this  more  and  more,  and  so 
compelling  the  secretions  into  the  stomachal  cavity,  constring- 
ing  the  glands  and  milking  them,  as  it  were,  into  the  food  to 
be  dissolved.  And  being  sandwiched,  so  to  speak,  between 
the  muscles  and  the  food,  the  open  ends  communicating  with 
the  latter,  this  action  in  the  muscles  could  not  otherwise  than 
have  that  effect  upon  the  glands.  It  must  do  so  in  the  very 
nature  of  things.  The  movements  which  set  in  in  the  stomach 
as  soon  as  food  is  introduced  for  disintegrating  the  boluses, 
would  have  the  effect,  then,  of  milking  the  secretions,  the 
object  being  to  bring  the  food  into  rapid  contact  with  the 
solvents.  But  this,  in  turn,  is  supplemented  by  the  force  in 
the  muscalaris  mucosa  for  increasing  the  action  in  the  tubes. 
Thus  numerous  small  fasciculi  proceeding  from  the  muscalaris 
mucosa,  extend  some  distance  up  the  tubes,  crossing  and 
interlacing  around  them,  and  so  forming,  as  it  were,  a  minute 
muscular  envelope  to  this  portion  of  the  follicles  for  expressing 
the  contents  into  the  stomachal  cavity.  Hence,  the  most  com- 
prehensive arrangements  obtain  in  the  stomach  for  compelling 
the  secretions  into  the  food. 

Finally,  since  the  gland  cells  are  contained  in  the  thinnest 
of  membranes  (membrana  proprii)  (Figs.  65,  66),  it  will  at 


PHYSIOLOGICAL    ANATOMY. 


199 


once  be  seen  that  mechanical  force  is  necessary  for  effecting 
rapid  expulsion  of  the  secretions. 
Concerning  the   Capillary  Network  in  the  Stomach. — In 


Fig.  65.— Vertical  Section  of  the  Human   Gastric  Mucous  Membrane,     a,  ridges  ;  6, 

peptic  glands. — Frey. 


Fig.  66.— Peptic  Glands  from  the  Human  Stomach  after  Treatment  with  Alkalies. — 

Frey. 

order  to  maintain  secretion  in  correspondence  with  the  amount 
of  ingesta  it  calls  for  appropriate  arrangements  in  the  capillary 
network  to  the  gastric  glands,  for  increasing  the  blood  supply, 


200  PHYSIOLOGICAL    ANATOMY. 

which  ebbs  and  flows  with  digestion,  and  in  correspondence 
with  this  being  increased  the  moment  that  food  is  introduced, 
diminishing  as  it  passes  out  of  the  cavity,  and  lowest  during 
the  interim. 

The  vascular  turgescence  which  the  stimulus  of  the  food  pro- 
duces in  the  gastric  mucous  membrane  (previously  pale)  has 
forcible  illustration  in  injected  preparations  (Fig.  07,  A  B) ; 
but  in  order  to  fully  understand  the  mechanics,  it  will  be 
necessary  to  go  a  little  deeper  in  the  tissues  passing  through 
and  through  the  membrane,  for  the  purpose  of  getting  at  the 
network  between  the  tubes  as  well,  as  also  to  obtain  a  view  of 
the  relations  which  they  sustain  to  the  arterial  and  venous 
systems,  or  the  afferent  and  efferent  vessels  to  the  mucous 
membrane,  which  is  given  us  in  the  following  beautiful  cut 
(Fig.  Gs)  by  that  accomplished  German  histologist  at  Zurich, 
from  whom  we  have  borrowed  so  copiously.  For  example,  it 
will  be  seen  that  the  arterial  feeder  (A),  in  passing  up  between 
the  tubes  to  reach  the  mucous  surface,  breaks  up  into  a  capil- 
lary network  for  feeding  the  gland  cells  in  the  tubes,  continu- 
ing thence  to  the  mucous  surface,  where  each  tubular  orifice  is 
surrounded  by  a  vascular  collar,  while  out  of  this  telangiectatic 
plexus  a  great  efferent  vessel  (d)  carries  the  blood  into  the 
portal  vein.  This  bird's-eye  view  will  give  some  idea  of  the 
vascular  arrangements  over  the  entire  surface  of  the  gastric 
mucous  membrane. 

Now,  then,  we  can  readily  understand  how  stimulation  of 
the  mucous  surface  by  springing  open  the  arterial  feeder 
under  the  action  of  the  special  nervous  ganglia  should  at  once 
produce  a  rush  of  blood  into  the  plexuses  ;  the  high  pressure 
in  the  arterial  system  compelling  this  to  be  done  the  instant 
that  expansion  sets  in,  and  which,  of  course,  should  increase 
the  action  in  the  gland  cells  correspondingly  for  maintaining 
'  this  in  due  proportion  with  the  quantity  of  ingesta.  But  the 
circumstance  to  which  attention  is  directed  more  especially  is 
the  great  relative  size  of  the  efferent  vessel  to  the  plexuses 
(d),  and  which  is  certainly  needing  explanation,  since  the 
amount  of  depletion  to  which  the  long-meshed  capillaries  are 
necessarily  subject  for  supplying  the  gland  cells  before  reach- 
ing the  round-meshed  network  upon  the  surface  would  natur- 


PHYSIOLOGICAL    ANATOMY. 


201 


ally  lead  one  to  expect  the  very  opposite  of  this,  or  a  small 
rather  than  a  large  efferent  vessel.  Nevertheless,  we  have 
this  great  vessel  springing  out  of  this  last  network ;  and 
since  this  can  effect  no  action  in  the  peptic  glands — being  at 
the  free  end  of  the  tubes — it  must  have  reference  to  other 
important  functions,  either  absorption,  or  else  to  pour  out  gas 
in  the  intestine  ;  most  probably  both  are  included.  It  will  thus 
be  seen  from  the  arrangements  that  obtain  in  the  parts,  that 
rapid  absorption  is  made  inevitable  from  the  very  nature  of 
things,  dialyzable  with  nou-dialyzable  substances,  otherwise 
inexplicable ;  notably,  the  rapid  absorption  of  alcohol, 
which  should  never  enter  the  circulation  at  all,  never- 
theless is  rapidly  absorbed  ;*   since  the  high  pressure,  which 


Fig.  67.— Appearance  of  the  Lining  Membrane  of  the  Stomach,  in  an  Injected  Prepara- 
tion. A,  from  the  convex  surface  of  the  rugae  ;  B,  from  the  neighborhood  of  the 
pylorus,  where  the  orifices  of  the  gastric  follicles  occupy  the  interspaces  of  the  deepest 
portions  of  the  vascular  network. — Carpenter. 

is  produced  in  the  stomach  by  means  of  the  firm  con- 
traction in  the  circular  muscles  must  inevitably  have 
this  effect  upon  the  liquid  contents,  compelling  them 
through  the  membranes.  Furthermore,  the  necessity  for  such 
short  and  expeditious  journey  of  the  fluids  to  the  blood,  is 
made  sufficiently  obvious  in  the  case  of  water,  for  the 
enormous  demands  in  the  nutritive  processes  and  for  main- 
taining a  balance  in  body-temperature,  which  is  accomplished 

*  "The  rapidity  with  which  alcohol  is  absorbed  in  the  stomach  is  forcibly 
shown  by  the  experiments  of  Dr.  Percy,  who  found  that  when  strong  alcohol 
was  injected  into  the  stomach  of  dogs,  the  animals  would  sometimes  fall  in- 
sensible to  the  ground  immediately  upon  the  completion  of  the  injection,  their 
respiratory  and  cardiac  movements  ceasing  within  two  minutes  ;  and  that  on 
post-mortem  examination  in  such  cases,  the  stomach  was  nearly  empty,  while 
the  blood  was  highly  charged  with  alcohol."  "Experimental  Inquiry 
Concerning  the  Presence  of  Alcohol  in  the  Ventricles  of  the  Brain,"  p.  61. 


202 


PHYSIOLOGICAL    ANATOMY. 


by  surface  evaporation,  would  soon  make  it  impossible  to  carry 
on  circulation  ;  hence,  this  open  door  for  rapidly  filling  the 
vessels  and  the  eagerness  with  which  thirst  is  quenched.  So, 
then,  this  circumstance,  in  the  special  anatomy  of  the  organ, 
is  strictly  in  accord  with  the  physiological  requirements,  the 
results  of  daily  experience  and  physiological  experiment. 
Of  course,  absorption  is  in  correspondence  with  the  stimulus 
to  action,  and  alcohol  being  a  powerful  stimulant,  producing 
rapid  expansion  in  the  vessels,  with  energetic  action  in  the 


Fig.  68.— Vascular  Network  of  the  Human  Gastric  Mucous  Membrane — half  diagram- 
matic.— Frey.  A,  fiue  arterial  twig,  which  breaks  up  into  a  long-meshed  capillary 
network  (b),  which  passes  again  into  a  round-meshed  one  (c)  around  the  openings  of 
the  glands.     From  this  latter  the  vein  (the  large  dark  vessel)  takes  its  origin. 

muscular  walls,  which  contract  firmly  upon  it,  the  most 
favorable  conditions  would  obtain  for  compelling  rapid  absorp- 
tion, the  great  tenuity  of  the  liquid  also  favoring  it.  Indeed, 
every  one  knows  the  liquids  are  rapidly  absorbed  in  the 
stomach,  while  the  mechanics  is  sufficiently  obvious,  the  force 
in  the  muscular  walls  being  available  for  this  purpose. 
Finally,  another  important  fact  remains  for  mention,  notably 
the  air  secreted  in  the  cavity,  passing  out  through  the  surface 
capillaries ;  a  quantity  of  air  being  also  ingested  with  the 
food,  and  which  assists  in  distending  or  ballooning  the  organ, 


SECRETION  of  gases  in  the  stomach.  203 

smoothing  out  the  folds  in  the  mucous  membrane,  and  making 
the  force  in  the  walls  available  in  the  work  of  digestion  and 
absorption  ;  at  the  same  time  acting  mechanically  for  effecting 
rapid  disintegration  of  the  boluses.  The  oxygen  carried  in 
with  the  atmosphere  passes  into  the  blood  through  the 
plexuses — a  remnant  of  the  old  or  primitive  method  of  respi- 
ration ;  while  carbonic  acid  passes  out  of  the  blood  into  the 
stomach  in  large  quantities,  for  increasing  pressure  in  the 
organ,  the  force  in  the  arterial  system  at  the  same  time  aiding 
it.  And  not  carbonic  acid  only,  but  nitrogen  as  well,  the 
two  gases  being  thus  poured  into  the  stomach  and  the  other 
portions  of  the  intestinal  canal  from  the  surface  capillaries 
during  digestion,  the  mechanics  being  fundamentally  the  same 
throughout,  and  which  is  regulated  by  nervous  force  so  as  to 
maintain  a  given  amount  of  pressure  in  the  canal  and  the 
cavity  of  the  abdomen.  Moreover,  it  would  explain  the  fol- 
lowing well-known  facts,  otherwise  inexplicable  ;  notably:  1st. 
The  function  possessed  by  the  intestines  of  rapidly  secreting 
air,  which  is  proven  to  demonstration  by  drawing  out  a 
loop  of  intestine,  isolating  it  by  ligature,  freeing  it  of  its 
contents,  and  returning  it  again  to  the  abdomen,  the  collapsed 
"walls  soon  becoming  distended,  as  previously. 

2d.  It  would  account  for  the  disappearance  of  carbonic  acid 
and  nitrogen  in  respired  air.  For  example :  the  carbonic  acid 
expired  does  not  represent  the  chemical  equivalent  of  the 
oxygen  inspired  ;  also,  that  an  amount  of  nitrogen  disappears ; 
while  both  are  contained  in  arterial  blood  and  are  present  in 
large  quantities  in  the  intestines,  especially  during  digestion, 
which  they  serve  to  expedite  by  their  mechanical  action,  and 
increase  the  absorptive  processes  correspondingly,  as  before  re- 
marked. 

Finally,  it  would  explain  the  rapid  evolution  of  gas  in  the 
intestines  produced  by  a  vegetable  diet,  which  is  harder  to 
digest  than  one  of  meats  ;  also,  why  the  condition  of  colic  can 
be  induced  by  substances  that  resist  digestion,  and  which 
stimulate  an  over-production  of  the  gases,  leading  to  distension 
and  pain.  And  that  this  is  not  due  to  fermentative  processes, 
is  proven  by  the  fact  that  the  gases,  when  liberated — as  I  have 
done  by  means  of  a  trocar  and  canula  in  the  horse — are  perfectly 


204  IMPORT    OF   THE   MOVEMENTS    IN   DIGESTION. 

free  from  any  odor  of  putrefaction  or  fermentation ;  in  fact, 
nearly  odorless. 

Also,  the  rapid  evolution  of  the  gases  in  dyspepsia,  hys- 
teria, etc.,  in  "which  they  are  poured  out  with  extraordinary 
rapidity,  producing  enormous  distensions  of  the  abdomen  in 
the  shortest  space  of  time. 

Furthermore,  the  power  to  secrete  air  to  subserve  other  uses 
than  those  immediately  connected  with  the  digestive  and 
absorptive  processes  is  seen  in  the  air-bladders  of  fishes, 
by  means  of  which  they  buoy  themselves  in  the  media,  raising 
and  lowering  the  body  readily  to  any  depth  by  simply  expand- 
ing the  air-sacs  and  body-walls,  thereby  diminishing  or  increas- 
ing body-density,  according  to  whether  they  desire  to  ascend  or 
descend,  as  the  case  may  be  ;  at  the  same  time,  they  expedite 
it  by  the  action  in  the  fins  and  tail.  Another  application  of 
the  principle  is  seen  in  the  eggs  of  birds,  etc.,  in  the  formation 
of  an  air-charnber  (Fig.  141),  by  means  of  which  the  rhythmi- 
cal changes  in  pressure  can  be  made  within  the  egg.  in  connec- 
tion with  the  actions  in  the  heart,  etc  ,  otherwise  impossible. 
Finally,  we  have  this  latest  application  in  the  intestines  for 
expediting  the  digestive  and  absorptive  processes  in  the  mam- 
malia, of  which  we  have  been  speaking. 

Concerning  the  Movements  in,  the  Stomach. — According  to 
Beaumont,  soon  after  digestion  sets  in  the  stomach  becomes 
divided  by  a  circular  constriction  into  two  distinct  compart- 
ments, in  which  the  action  differs;  the  rhythmical  contractions 
and  expansions  in  the  cardiac  end  being  slow  and  gentle,  while 
in  the  pyloric  they  are  more  rapid  and  energetic. 

This  division  in  the  organ  is  produced  by  a  firm  contraction 
of  transverse  muscular  fibres  near  the  pyloric  end,  the  smaller 
compartment  occupying  about  one-third  and  the  large  about 
two-thirds  of  the  organ  ;  while  between  the  two  ends  a  con- 
stant current  of  the  fluids  is  maintained,  by  means  of  which 
the  boluses  are  soon  disintegrated  and  penetrated  in  every 
portion  with  the  gastric  juices.  "As  the  alimentary  bolus 
enters  the  stomach  by  the  cardiac  opening,  it  turns  to  the  left, 
descends  into  the  great  pouch,  and  follows  the  great  curvature 
to  the  pyloric  end.  It  then  returns  to  the  cardiac  orifice  by  the 
lesser  curvature,  and  takes  again  the  same  course  as  before. 


IMPOKT   OF   THE   MOVEMENTS   IN  DIGESTION.  205 

While  these  revolutions,  so  to  speak,  of  the  alimentary  mass 
are  going  on,  the  food  is  turned  over  and  over,  so  that  it  be- 
comes intimately  mixed  with  the  digestive  fluids  and  subjected 
to  a  certain  amount  of  trituration."  "  When  the  thermometer 
bulb  arrived  at  the  contracted  septum,  which  was  three  or  four 
inches  from  the  pyloric  end,  it  was  at  first  stopped  by  the  forci- 
ble contraction  ;  but  in  a  short  time  there  was  a  gentle  relaxa- 
tion, which  allowed  it  to  pass,  when  it  was  drawn  quite  forcibly 
for  three  or  four  inches  toward  the  pyloric  opening  When 
in  this  portion  of  the  stomach,  the  bulb  was  firmly  grasped 
and  made  to  undergo  a  spiral  motion  ;  and  if  drawn  forcibly 
out,  it  gave  to  the  fingers  the  sensation  of  being  held  by  a 
strong  suction  force.  As  soon  as  relaxation  occurs,  the  bulb 
is  passed  back  to  the  seat  of  stricture,  and,  when  pulled 
through  this,  it  moves  freely  in  the  great  cavity  Each  of 
these  revolutions  occupied  from  one  to  three  minutes.  They 
were  slower  at  first  than  after  digestion  had  been  somewhat 
advanced. ' ' 

These  graphic  excerpts  of  practical  observations  made  upon 
the  stomach  during  digestion  would  leave  no  doubt,  then,  of 
the  mechanical  principle  which  applies  for  increasing  diges- 
tion and  absorption,  namely,  rapid  rhythm'' cal  changes  in 
pressure,  the  alternating  contractions  and  expansions  in  the 
two  ends  serving  to  maintain  a  current  of  the  viscid  chyme 
through  the  limits  in  the  organ,  but  all  the  while  flowing  from, 
high  to  low  pressure,  in  conformity  with  organic  law.  When 
contraction  sets  in,  however,  the  sudden  reversal  of  the  cur- 
rent which  this  effects  produces  the  spiral  motion — a  minia- 
ture whirlpool,  as  it  were — while  the  suction-force  spoken  of 
was  simply  a  pulling-force  produced  by  the  current  setting 
into  the  compartment  and  the  energy  of  the  contraction  twist- 
ing the  current  upon  itself,  and,  of  course,  pulling  the  bulb 
along  with  it.  As  soon  as  expansion  is  effected,  however,  this 
restores  the  horizontal  line  again,  when  the  current,  deflected 
by  the  opposite  wall,  simply  carries  the  bulb  back  to  the  seat 
of  constriction.     It  is  all  simple  enough. 

When  the  bolus  enters  the  stomach,  it,  of  course,  falls  into 
the  surface  current  which  sets  into  the  great  cul  de  sac,  the 
force  in  the  muscular  contractions  in  the  pyloric  end,  by  reason: 


206 


MTTSCl  LAK   FORCE   IN  DIGESTION. 


of  greater  energy,  producing  this  surface  current ;  while  "the 
food  is  turned  over  and  over"  by  reason  of  the  deeper  cur- 
rent setting  in  the  opposite  direction  along  the  floor  of  the 
organ  into  which  the  boluses  project,  and,  of  course,  causing 
them  to  turn  over  and  over.  Furthermore,  it  is  also  easily 
perceived,  from  the  nature  of  the  mechanical  adjustments  that 
obtain  in  the  gastric  glands  and  capillaries,  that  secretion  and 
absorption  are  maintained  in  correspondence  with  the  c/iurn- 
ing-actlon,  in  the  stomach.  I  do  not  see  necessity  for  adding 
another  word,  the  matter  being  so  obvious.     Something  might 


Fig.  69. — External  Muscular  Fibres  in  the  Stomach. — Gray. 

be  said,  however,  in  regard  to  the  muscles  in  the  walls  of  the 
stomach,  which  is  the  real  force  in  digestion  and  absorption  ; 
as  it  is  through  them  the  rhythmical  changes  in  pressure  are 
produced.  They  are  nearly  all  circular  muscles  (Figs.  69,  70) ; 
with  these  exceptions,  however,  that  upon  the  external  surface 
scattering  longitudinal  fibres  extend  along  the  sides,  and 
greater  and  lesser  curvatures  (thicker  in  these  two  localities), 
with  a  thick  fan-shaped  layer  spread  out  over  the  upper 
portions  of  the  great  cul  de  sac  formed  by  the  longitu- 
dinal  muscles   in  the  oesophagus  (Fig.  69) ;  while   upon   the 


MUSCULAR  FOKCE  IN   DIGESTION. 


207 


internal  surface  we  have  a  wide,  thin  layer  of  longitudinal 
muscles,  embracing  the  great  cul  de  sac  and  extending  some 
distance  toward  the  pyloric  end,  but  fading  out.  however,  and 
disappearing  upon  the  sides  of  the  organ  (Fig.  70).  The  firm 
constriction  near  the  pyloric  end  is,  of  course,  produced  by 
the  contraction  of  a  band  of  circular  muscles,  as  are  also  the 
rhythmical  contractions  and  expansions  in  the  two  ends,  while 
i  he  longitudinal  fibres  in  the  great  cul  de  sac  should  greatly 
expedite  the  pumping  action  by  compelling  the  contents 
toward  the  pyloric  end  during  their  contraction,  since  it  must 

!l 


Pig.  70. — Fibres  seen  with  the  Stomach  hvcr^M. — oappey.  1,  LKsophagus  ;  2,  circular 
fibres  at  the  oesophageal  opening  :  3,  3.  circular  fibres  at  the  lesser  curvature  :  4,  4, 
circular  fibres  at  the  pylorus  ;  5,  5,  6,  7.  8.  oblique  fibres  :  9,  10,  fibres  of  this  layer 
covering  the  greater  pouch  ;  11,  portion  of  the  stomach  from  which  these  fibres  have 
been  removed  to  show  the  subjacent  circular  fibres. 

inevitably  effect  a  corresponding  shortening  in  the  longi- 
tudinal axis  of  the  organ.  One  result  of  the  circular  con- 
striction would  be  to  shorten  the  longitudinal  fibres  embracing 
the  cul  de  sac  (Pig.  ?> ,  5,  6,  7,  h) ;  at  the  same  time,  by  giving 
them  a  firm  point  to  contract  upon,  would  make  their  action 
upon  the  cul  de  sac  still  more  effective  for  compelling  the 
contents  toward  the  pyloric  end,  while  the  greater  number  of 
the  circular  muscles  in  this  end,  together  with  the  diminished 
area,  should  increase  the  energy  in  the  rhythmical  expansions 
and  contractions  correspondingly,  increasing  the  frequency  and 


208  ACTION    IN  THE  SMALL   INTESTINES. 

force  of  the  pumping  actions  in  this  end  for  compelling  move- 
ment in  the  contents. 

In  this  manner,  then,  muscular  force  is  so  distributed  as  to 
make  it  effective  upon  every  portion  of  the  gastric  contents 
for  compelling  rapid  digestion  and  absorption,  the  energy 
of  whicli  is,  of  course,  regulated  by  nervous  force,  while  this 
in  turn  is  determined  by  the  requirements  in  the  organism. 
Furthermore,  we  can  also  readily  understand  how  the  mechan- 
ical action  in  the  air-cushion  should  increase  the  churning? 
action  correspondingly,  since  the  immediate  result  is  a  great 
gain  in  srjace  in  which  to  effect  it ;  at  the  same  time  the  air  serves 
for  disintegrating  the  boluses  by  transmitting  the  force  in  the 
muscular  waits  upon  them.  In  fine,  the  air  is  a  great  and 
essential  factor  in  the  mechanics  in  the  abdomen. 

Concerning  the  Action  in  the  Small  Intestines. — The  mu- 
cous membrane  of  the  small  intestines  is  more  complicated 
than  that  of  the  stomach  ;  but  we  shall  pass  over  the  more 
prominent  features  as  rapidly  as  we  can,  for  the  purpose 
of  showing  the  principle  in  the  mechanics  and  the  great  role 
which  is  performed  by  the  muscular  cylinder.  Beginning  with 
the  duodenum,  we  have  the  mucous  membrane  arranged  in 
crescentic  folds  [valvulo3  conniventes]  (Fig.  SG),  which  continue 
throughout  the  jejunum  and  into  the  ileum  to  about  the  middle 
portions,  where  they  fade  out  and  disappear.  They  extend 
about  one-half  to  three-fourths  around  the  cylinder,  springing 
from  every  portion  of  the  circumference,  the  evident  object 
being  for  increasing  the  secretory  and  absorptive  processes,  at 
the  same  time  providing  for  free  action  in  the  muscles  or 
peristalsis  ;  but  in  contraction  they  are  made  more  prominent. 
Through  the  rest  of  the  ileum  it  is  smooth,  and  in  the  large 
intestine  is  evenly  applied  over  the  surface  of  the  sacculi ; 
while  in  the  rectum  it  is  again  thrown  into  longitudinal  folds, 
as  in  the  oesophagus  and  stomach,  as  the  necessary  provision 
for  effecting  the  wide  expansions  which  occur  in  this  pouch, 
which  functions  as  a  receptacle  for  the  faeces  before  final  ex- 
pulsion. The  three  valvular  folds  corresponding  with  the 
upper,  middle  and  vesicle  portions  tend  to  delay  the  onward 
movement,  as  in  the  sacculi  of  the  colon,  at  the  same  time 
increasing  the  absorptive  surface  ;  while  the  arrangement  that 


PHYSIOLOGICAL    ANATOMY. 


209 


obtains  at  the  ileocsecal  valve  serves  for  obviating  reflux  in 
the  small  intestines. 

The  following  diagram  will  give  some  idea  of  the  arrange- 
ment of  the  glandular  structures  in  the  duodenum  (Fig.  7 J). 
In  the  so-called  Brunnef  s  glands  (c)  found  in  the  duodenum 
only — in  the  upper  portions  especially — we  have  a  group  of 
open  gland  vesicles  of  microscopic  dimensions,  flask-shaped, 
with  short  necks,  that  discharge  their  contents  through  an 
axial  canal ;  and  are  simply  transitional  forms  of  the  simple 
gland  tubes  that  have  coalesced  near  the  blind  ends,  forming  a 
new  anatomical  unit,  or  a  so-called  racemose  gland  (Frey). 
In  the  compound  peptic  glands  the  same  circumstance  is  seen, 
only  that  coalescence  occurs  higher  up  the  tubes  and  nearer 


Fig.  71.— Vertical  Section  of  Mucous  Membrane  of  Duodenum,  shoTring  Brunner's 
glands,  a,  Follicles  of  Lieberkiilin  ;  b,  cellular  coat  of  intestine  ;  c,  Brunner's  glands  ; 
d,  annular  fibres  of  muscular  coat ;  e,  longitudinal  fibres  of  muscular  coat. — Wilson. 

the  mucous  surface,  is  also  limited  to  several  tubes  only  ;  here 
a  whole  group  coalesce. 

They  secrete  a  clear  alkaline  mucus,  free  of  form  elements. 
Arranged  around  the  central  canal  as  foot-stalk,  they  bear 
striking  resemblance  to  a  bunch  of  grapes  (Fig.  72).  The 
relations  they  sustain  to  the  bases  of  the  villi  is  shown  in  the 
following  cut  (Fig.  73).  The  simpler  forms,  however,  are  seen 
in  the  crypts  or  follicles  of  LieberJcuJin  (Fig.  71,  a),  arranged 
perpendicularly  to  the  surface,  upon  which  they  open  by  circular 
apertures,  and  occupy  the  whole  free  surface  of  the  small  and 
large  intestines,  opening  around  the  bases  of  the  villi  and  the 
solitary  glands.  Their  walls,  however,  are  exceedingly  thin, 
at  times  nearlv  indistinguishable  from  the  surrounding  tissues 


2J0 


PHYSIOLOGICAL   ANATOMY. 


(Fig.  74).  The  contents  of  these  crypts,  unlike  those  of 
Bru aner" s  glands,  consist  of  delicate,  columnar,  nucleated 
cells  (a),  with  the  bases  resting  against  the  membrana  propria, 


Fig  72. — One  of  Brumwr'.-:  Racemose  '.  Hands,  from  the  Human  Being. — Prey. 


Fig.  73. — Brunner'ls  Glands,  from  the  Duodenum. — Frey.     a,  Villi  ;  b,  bodies  of  glands  ; 
c,  excretory  canal  opening  between  the  villi. 

the  apices  presenting  in  the  crypt  (Fig.  74,  K ;  Fig.  75,  d). 
These,  together  with  the  axial  canal,  may  be  seen  in  every  trans- 
verse section  (Fig.  75,  d).    According  to  Schulze,  between  these 


PHYSIOLOGICAL    ANATOMY. 


211 


cells  other  goblet  cells  may  present  themselves  (Fig.  76,  a). 
They  were  thought  to  be  confined  to  the  surface  of  the  villi. 
While  the  orifices  of  the  Lieberkiihnian  follicles  are  neces- 


Fig.  74.— Follicle  of  Lieberkuhn,  greatly  magnified. — Verson.  K,  follicle  ;  a,  a,  epithe- 
lium ;  d,  adenoid  tissue,  from  which  the  cells  have  been  removed  by  penciling  ;  T, 
fibrous  tissue  on  the  opposite  side. 


Fig.  75.— From  the  Small  Intestine  of  the  Rabbit;  a,  Tissue  of  the  mucous  membrane  ; 
6,  lymphatic  canal ;  c,  an  empty  transverse  section  of  a  gland  of  Lieberkuhn  ;  d, 
another  of  the  same  occupied  by  cells. — Frey . 


212  PHYSIOLOGICAL    ANA  I  OMY. 

sarily  separated  by  the  villi,  the  tubes  dilate  beneath  them  in 
such  manner  as  almost  to  bring  them  in  contact,  leaving  only 
small  interspaces  for  the  passage  of  vessels  and  muscular 
fasciculi.  Finally,  we  have  to  mention  the  lymphoid  follicles 
or  glandules  solitaries,  that  are  scattered  over  the  mucous 
membrane  of  the  small  intestine,  and  when  a  gminated  forming 
Peyer's  patches,  and  which  are  analogous  to  lymphatic 
glands.  They  are  situated  either  in  the  tissue  of  the  mucous 
membrane,  or,  when  of  considerable  length,  project  down  into 
the  submucosa,  as  occurs  in  Peyer's  patches,  in  which  the 
bases  of  the  follicles  are  approximated  to  the  circular  mus- 
cles, while  the  apices  project  upon  the  mucous  surface  sur- 
rounded by  villi,  the  fibres  of  the  muscalaris  mucoscs  being 
separated  and  pushed  aside  by  the  enlarged  follicles  (Pig.  77, 
m,    S,  K).     They  are  ductless  glands,   and  the  means  for 


Fig.  76. — Epithelial  Cells  from  a  Human  Intestinal  Villus  (after  Schulze).    a,  Goblet 
cells  ;  b,  ordinary  elements. 

increasing  circulation  in  them  and  for  compelling  out  the  secre- 
tions possesses  special  interest. 

They  occur  opposite  to  the  attachment  of  the  mesentery, 
consequently  their  caps  are  kept  constantly  submerged  in  the 
intestinal  fluids,  gravitation  compelling  this  circumstance  ; 
are  from  one  to  three  inches  in  length  and  about  an  inch  in 
breadth,  are  covered  with  villi,  and  the  Lieberkiihnian  fol- 
licles form  a  circle  around  them  ;  are  more  numerous  in  the 
lower  portions  of  the  ileum,  while  they  vary  from  twenty  to 
thirty,  and  even  more,  in  number,  becoming  less  and  less, 
however,  as  age  advances.  The  usual  number  of  follicles  in 
a  patch  is  from  twenty  to  thirty,  but  they  vary  from  as  low 
as  three  to  seven,  while  the  large  patches  may  contain  from 
fifty  to  sixty  (Prey).  The  following  illustration  will  show  the 
character  of  the  mucous  membrane  in  the  large  intestine,  and 
the  relations   it   sustains   to   the  circular  muscles  (Fig.    7). 


PHYSIOLOGICAL    ANATOMY. 


213 


The  crypts  of  Lieberkulm  continue,  but  the  villi  extend  no 
further  than  the  free  border  of  the  ileo-csecal  valve.  It  pre- 
sents the  same  structure  and  arrangement  of  its  constituent 
parts  as  the  small  intestine,  of  which  it  is  a  direct  continua- 
tion. Now,  then,  the  point  to  which  attention  is  specially 
directed  concerns  the  action  in  the  muscular  cylinder  in  which 
the  mucous  membrane  is  placed.    From  the  nature  of  these 

z 


Fig.  77.— A  Longitudinal  Section  of  the  Small  Intestine  of  a  Rabbit,  through  a  Peyer's 
Patch  (S).— Verson.  Z,  Z,  villi  ;  J,  follicles  of  Lieberkuhn  ;  K,  cap  of  a  gland  ;  m, 
muscalaris  mucosae  ;  C,  C,  submucosa  ;  S,  glands  of  Peyer  ;  B,  circular  muscles  ;  L, 
longitudinal  muscles  ;  P,  peritoneum. 

anatomical  dispositions  in  the  mucous  membrane,  and  the  me- 
chanical adjustments  that  obtain  in  the  intestine,  it  follows 
that  contraction  in  the  muscles  must  have  the  effect  of  forcibly 
compressing  the  gland  follicles  and  lymphoid  follicles  (Figs. 
71-78),  and  compelling  the  secretions  out  of  them,  gently  but 
effectually  constringing  them,  and  milking  them,  so  to  speak, 


214 


PHYSIOLOGICAL  ANATOMY. 


of  their  contents  by  means  of  the  uniform  compression  ex- 
erted through  the  elastic  air-cushion  under  the  energy  in  the 
muscles. 

Sandwiched  as  they  are  between  these  forces,  with  the  mus- 
cles as  the  firm  floor  of  support,  the  cushion  pressing  against 
them,  it  could  not  otherwise  than  have  this  effect  upon  the 
glands.  It  at  once  settles  the  matter.  The  air  is  in  there  and 
it  cannot  get  away,  and  the  muscles  certainly  contract;  the 
rest  follows  as  an  inevitable  sequence.  Xow,  then,  to  this 
must  be  added  the  tine  adjustments  in  the  muscularis  mucosae, 


Fig.  78. — Section  of  the  Large  Intestine  of  a  Rabbit.  J,  crypts  of  Lieberkiihn ;  a, 
epithelium  ;  b,  mucosa  ;  m,  muscularis  mucos.-e  :  s,  submucosa  ;  R,  circular  muscular 
layer  ;  L,  longitudinal  muscular  layer  ;  p,  peritoneum. — Verson. 

and  the  thing  will  be  complete.  It  must  be  borne  in  mind  that 
the  viscid  secretions  are  difficult  to  move  ;  hence,  force  must  be 
applied  directly  to  the  follicles,  which  is  done  by  means  of 
small  muscular  fasciculi  extending  from  t lie  muscularis  mu- 
cosas between  the  acini  of  Brunners  glands  and  between  the 
follicles  of  Lieberkuhn  ;  but  are  much  more  numerous  in  the 
villi,  running  upward  into  the  parenchyma,  so  as  to  form  the 
wall  of  the  central  lacteal  (Fig.  98,  m,  I). 

This,  together  with  the  force  in  the  circular  muscles,  exerted 
in  the  manner  as  stated,  compel  Qe  secretions  out  of  the  fol- 


PHYSIOLOGICAL  ANATOMY.  215 

licles.  It  now  remains  to  take  np  the  work  of  absorption, 
for  which  these  adjustments  are  equally  effective,  bringing 
out,  however,  some  fresh  features  in  the  mechanics  which  are 
extremely  beautiful  and  charming  to  look  upon.  There  is 
nothing  more  wonderful  than  animal  mechanics. 

The  following  illustration  will  give  some  idea  of  the  general 
appearance  of  the  mucous  membrane  of  the  small  intestines 
(Fig.  79).  By  means  of  the  villosities,  together  with  the  cres- 
cendo folds  in  the  membrane,  the  mucous  surface  is  enor- 
mously increased  ;  but  the  former  have  more  to  do  with  the 
absorptive  processes  simply,  though,  by  reason  of  the  special 


Fig.  79. — Portion  of  the  Mucous  Membrane  of  Small  Intestines,  showing  the  villi  and  the 
apertures  of  the  simple  follicles,  magnified  19  times. — Wilson.  In  the  hollows 
between  the  villi  are  seen  the  apertures  of  simple  f  ollieles  (b)  ;  and  near  the  bottom 
of  the  figure  is  a  zone  of  follicles  (a)  surrounding  a  solitary  gland. 

capillary  arrangements  which  obtain  around  the  outlets  to  the 
follicles  of  Lieberkiihn  (Fig.  80,  d),  or  the  same  as  obtains  in 
the  stomach  (Fig.  68,  c),  absorption  goes  on  quite  actively, 
independently  of  the  villi ;  only  that  it  is  not  so  rapid  as  in 
the  stomach,  for  the  vascular  network  is  not  so  extensive, 
the  greater  portion  of  the  capillaries  going  into  the  villi 
And  it  will  be  seen  that  here  also  there  is  a  great  contrast 
in  the  size  of  the  arterial  feeders  and  the  discharging  vessels 
or  veins  (a,  c),  the  veins  being  out  of  proportion  to  the  arterial 
feeders,  the  import  of  which  has  already  been  stated.  Now, 
then,  between  these  vascular  loops  and  the  liquids  to  be  ab- 
sorbed is  the  layer  of  columnar  epithelium  ;  and  the  question 


216 


PHYSIOLOGICAL  ANA'IOMY 


before  us  concerns  the  manner  tins  barrier  is  passed,  or  bow 
fat,  albumen  and  other  non-dialyzable  substances  effect  a  rapid 
passage  into  the  interior  of  the  cells,  the  membrane  (membrana 
propria)  beyond,  and  so  get  into  the  interior  of  the  villi  to  the 
capillaries  and  central  lacteal.  Furthermore,  keep  the  fact  in 
mind  that  the  fat  is  simply  emulsified,  not  saponified,  while 
albumen  passes  in  freely  unchanged,  especially  when  given  in 
large  quantities,  thereby  showing  conclusively  the  existence 
of  a  mechanical  force  for  compelling  them  through  the 
membranes ;    otherwise  is  inexplicable.     This   mechanics  is 


Fig.  80. — Vascular  System  of  an  Intestinal  Villus  in  the  Rabbit,  a,  The  arteries 
(shaded),  breaking  up  first  into  a  capillary  network  around  the  glands  of  Lieberkuhn 
(d)  ;  b,  network  of  capillaries  in  the  villus  ;  c,  venous  vessels  (unshaded). — Frey. 

also  easily  understood.  Thus,  the  columnar  epithelia  are 
covered  by  a  perforated  lid  of  thickened  epithelium 
(Fig.  bl,  a,  b),  which  acts  as  a  sieve  or  colander  for  strain- 
ing the  chyle,  while  the  muscular  cylinder  contracts 
for  forcing  it  into  the  epithelia.  whence  it  is  passed  in  due 
time  into  the  parenchyma  of  the  villi,  and  compelled  thence 
into  the  capillaries  and  the  central  lacteal,  the  coarser  par- 
ticles passing  by  the  vessels  into  the  latter,  which  functions 
as  a  drainage  system  to  the  villi.  The  lids  fit  neatly  over  the 
surface  of  the  cells,  and  being  in  close  apposition  and  firmly 
consolidated  by  animal  cement,  which  fills  up  the  interstices 


PHYSIOLOGICAL  ANATOMY.  217 

(Fig  82,  5),  present  an  appearance  not  unlike  a  mosaic  pave- 
ment (81,  b).  But  when  macerated  in  water,  they  are  easily 
loosened  by  pressure  (Fig.  <"■  2,  a) ;  the  splitting  up  of  the  lids 
through  the  perforations  giving  them  the  appearance  of  cilia, 
is  a  result  of  post-mortem  changes  in  the  cells.  Thus  con- 
stituted, then,  the  force  in  the  muscular  cylinder  is  readily 
brought  to  bear  for  compelling  the  liquid  aliment  into  those 
myriads  of  little  organs  (each  one  of  which  may  be  regarded 
as  a  distinct  gland  in  itself)  that  are  spread  out  like  a  sheet 
over  the  external  surface  of  the  villi  for  effecting  further 
metamorphosis  upon  the  aliment,  thence  into  the  parenchyma  of 
the  villi,  thence  into  the  vessels,  as  before  stated  ;  the  structures 
in  the  villus — epithelium,  blood-capillaries,  and  central  lacteal 
embraced  by  its  special  muscles  (Fig.  98,  Z,  m\  all  acting  to- 
gether and  in  harmonious  concert  with  the  force  in  the  muscu- 


Fig.  81. — Columnar  Cells  from  the  Small  Intestine  of  the  Rabbit,  a,  Side  view  of  cells 
with  thickened  raised  lids  traversed  by  pores  ;  6,  view  from  above,  in  which  the 
orifices  of  the  pores  appear  like  dots. — Frey. 

lar  cylinder  for  compelling  rapid  absorption.  This  additional 
force  in  the  fine  adjustment  spoken  of  in  the  muscularis 
mucosae  must  necessarily  increase  the  action,  since  the 
rhythmical  contractions  and  expansions  in  the  muscular 
fasciculi  that  penetrate  the  parenchyma  and  form  the 
wall  of  the  central  lacteal,  could  not  otherwise  than  have 
that  effect,  must  do  so,  in  the  very  nature  of  things,  acting 
as  a  little  suction-pump  and  increasing  circulation  in  the 
vessels  correspondingly.  And  with  high  intra-intestinal 
pressure  produced  by  the  muscular  cylinder  and  the  gases  in 
the  intestines,  together  with  this  pumping  action  in  the  villi, 
absorption  should  go  on  very  rapidly,  as  a  matter  of  course  ; 
while  the  rhythmical  expansions  and  contractions  taking  place 
in  the  gut  itself,  diffusing  the  aliment  over  the  mucous  sur- 
face, at  the  same  time  increasing  pressure  should  increase  the 
action  correspondingly.  And  through  these  combined  actions 
in  the  organ  effected  by  means  of  nervous  force,  we  can  very 


218 


RAFID   ABSORPTION   INEVITABLE. 


readily  understand  the  rapid  absorption  of  non-dialyzable 
substances,  otherwise  inexplicable.  Nay,  bow  even  particles 
of  finely-divided  charcoal  may  present  in  the  mesenteric  veins 
(Oesterlen);  and  why  similar  results  have  been  obtained  by 
Eberhard,  Mensonides,  and  by  Donders,  not  only  with  char- 
coal, but  also  with  sulphur,  and  even  with  starch,  the  latter 
substance  being  at  once  detectable  in  the  blood  by  the  iodine 
test.  Furthermore,  there  can  be  no  doubt  but  that  they 
enter  the  circulation  through  the  epithelial  cells  of  the  villi, 
as  the  presence  of  psorosperms  in  the  interior  of  these  has 
been  distinctly  seen  (Klebs);  while  they  are  nearly  double 
their  size  soon  after  digestion  sets  in,  and  of  a  milky  appear- 


Eig.  82.— The  same  cells.  At  a,  the  border  is  loosened  by  water  and  slight  pressure  ;  b, 
natural  condition  ;  c.  a  portion  of  the  lid  destroyed  ;  d,  e,  /,  the  latter  is  resolved 
into  a  number  of  rod-like  or  prismatic  pieces,  by  maceration  in  water. — Frey. 

ance  from  the  quantity  of  fat  they  contain  ;  and  that  fat  enters 
the  blood-vessels  also,  as  well  as  the  central  lacteals,  is  proven 
by  the  opalescent  whiteness  of  the  superficial  capillary  net- 
work, and  the  quantity  of  fat  contained  in  the  portal  veins, 
fully  one-half  the  fat  being  taken  up  by  the  veins.* 

And  with  all  these  muscles  in  the  gut,  the  fine  with  the 
coarse  adjustments  that  obtain,  the  nerves  for  operating  them, 
together  with  the  air-cushion  as  the  lever  for  effecting  the 
changes  in  pressure  upon  the  contents,  no  reason  on  earth 
presents  why  absorption  should  not  be  very  rapid.  And  the 
miserable  device  which  comes  to  us  as  a  suggestion  by  a  promi- 
nent English  microscopist,  that  the  fat  finds  its  way  to  the 
central  lacteal  by  passing  between  these  cells,  cannot  be 
entertained  for  a  single  moment  even  ;  since  this  would  pass  it 


Zawilski,  Ludwig's  Arbeiten,  1876,  p.  147. 


EAPID   ABSORPTION  INEVITABLE.  219 

through  the  long  line  of  cement  binding  these  minnte  organs 
together,  and  should  undermine  the  very  foundations  of  this 
beautiful  structure ;    while  it  would  leave   unexplained  the 
rapid  absorption  of  fat  by  the  blood  vessels,  together  with 
the  entire  anatomy  in  the  gut  as  means  to  ends.      What ! 
The  chief  force-producer  burrowing  under  the  foundations  in 
order  to  get  into  the  building  ?    Nein  !  Niclit  zu  glauben  I 
2s  ever  under  the  canopy  of  heaven  but  by  the  public  way 
through  the  open  front-door — i.  e.,  the  sieve-like  openings  in 
the  epithelium — with  the  force  in  the  muscles,  the  fine  and 
coarse  adjustments  for  assisting  the  passage.    The  one  explains 
everything  ;  the  other  explains  nothing,  which  at  once  exposes 
it.     Besides,    we  positively  know  the    fat  enters    the    cells 
very  rapidly,    so    that   they    soon    become    twice    the    size 
in  the  empty  condition,  and  the  fat  is  easily  detectable  in 
the  milky  appearance  it  imparts  to  the  cells.     For  increas- 
ing  the   action   we   have   only  to. increase  pressure  in  the 
cavity  by  contracting  the    muscular  walls  of    the   gut,   at 
the  same  time  increasing  the  action  in  the  villi,  the  whole  per- 
forming as  a  single  organ  only  for  pumping  the  fluids  into  the 
vessels.     Hence  the  slow  and  sustained  contraction  which  fol- 
lows peristalsis.     And  with  the  anatomical  dispositions  which 
obtain  in  the    organs  and  the  mechanical  principle  which 
applies  for  increasing  circulation,  nu  reason   presents   why 
digestion  and  absorption  should  not  be  made  commensurate 
with  the  physiological  requirements.      This  explains  every- 
thing ;  whereas  the  old  method  explains  little  or  nothing — 
nay,  is  pitiful  in  its   utter  helplessness  and  power  to  help 
itself ;  vainly  groping  in  the  tissues  with  the  rnicroscoTje  in 
search  of  the  key  to  the  vexed  problem,  not  to  be  found  with 
the  microscope,  but  in  the  organic  law  itself  underlying  animal 
structure.     And  from  this  stand-point  the  whole  mechanics  in 
the  animal  circulation  is  at  once  made  intelligible,  from  the 
cell  itself  to  the  cell-empire,  the  multitudinous  arrangements 
in  the  structures  falling  readily  into  line  in  regular  order  and 
succession.     But  in  the  absence  of  this  law  of  pressure  and 
fluid  equilibrium,  which  is  being  thus  incessantly  invoked  in 
the  measure  of  the  physiological  requirements,  all  are  inex- 
plicable, all  is  chaos. 


CHAPTER  X. 

KESPIRATION  AND   THE   PORTAL   CIRCULATION 

Circulation  in  the  Liver  Dependent  upon  Respiration — Mechanics  in  the  Diaphragm  and 
Walls  of  the  Abdomen  Respecting  it — The  Mesentery  a  Soft,  Elastic  Cushion  for 
Effecting  Gentle  Compression  of  the  Liver-Substance,  under  the  Action  in  the  Dia- 
phragm and  Walls  of  the  Abdomen  during  Inspiration  for  Increasing  its  Circulation 
— Mode  of  Demonstrating  this  Circumstance — Effect  upon  the  Portal  Vessels  and 
Lower  Cava-System — Absence  of  Valves  in  the  Veins  within  the  Abdomen,  save  in 
the  Pelvic  Viscera  Only — Explanation  for  the  Latter  Circumstance — Physiological 
Anatomy  of  the  Liver,  with  Reference  to  Circulation — Why  the  Hepatic  Veins  are 
Incorporated  with  the  Liver-Substance — Relations  of  the  Portal  System  to  the 
Hepatic  Viens — Why  Rhythmical  Compression  of  the  Liver-Substance  during  Res- 
piration should  Increase  Circulation  in  the  Inter-  and  Intra-Lobular  Vessels  Corre 
spondingly — Automatic  Action  in  the  Portal  Vessels  Synchronous  with  Respiration 
for  further  Increasing  it — Mechanics  for  Circulating  Bile — The  Action  in  the  Gall- 
Bladder  an.l  Bile-Ducts — The  Action  in  the  Duodenum,  in  Connection  with  the 
Biliary  and  Pancreatic  Secretions — Adjustments  in  the  Viscera  Necessitated  by  the 
Action  in  the  Diaphragm — Mechanics  Connected  with  the  Openings  in  the  Dia- 
phragm— (Esophagus  Constricted  during  Inspiration,  while  the  Vena-Cava  Lumen 
is  thrown  Widely  Open  ;  as  also  the  Aortic — Elongation  and  Contraction  of 
CEsophagus  with  Inspiration  and  Expiration — Ditto  Venae  Cava? — Ditto  Portal 
Vessels  and  Renal  Veins — Similarity  in  the  Anatomical  Dispositions  of  the  Muscles 
in  these  Organs. 

Before  taking  up  the  nervous  apparatus  to  the  intestines,  it 
will  be  necessary  to  make  a  rapid  survey  of  the  glandular  ap- 
pendages and  the  incidental  adjustments  rendered  necessary 
by  the  action  in  the  diaphragm  ;  since  the  nervous  apparatus 
effects  coordination,  to  the  end  that  the  whole  should  work 
together  harmoniously,  without  let  or  hinderance,  within  cer- 
tain prescribed  limits,  that  automatism  may  be  maintained  in 
the  organs  and  structures,  which  is  necessary  to  the  existence 
of  the  organism.  It  is  very  comprehensive.  But  it  is  equally 
manifest  that  all  of  it  is  based  upon  pressure,  and  the  power 
of  producing  rapid  rhythmical  changes  in  pressure  for  in- 
creasing circulation  and  compelling  this  to  be  in  correspond- 
ence with  the  physiological  requirements,  which,  of  course, 
would  include  the  secretions  in  the  glands,  with  the  arrange- 


THE  ABDOMEN   A   VENTRICLE.  221 

ments  for  compelling  them  into  the  cavity  of  the  intestine.  For 
example,  we  can  readily  understand  why  the  liver- substance  is 
spread  out,  so  to  speak,  upon  the  incurvated  surface  of  the 
diaphragm,  to  which  it  is  fastened  by  the  overlying  peri- 
toneum (Fig.  25,  L),  and  why  it  is  itself  excavated  for  receiving 
the  abdominal  viscera  which  rest  against  its  surface,  since  this. 
would  favor  the  rhythmical  compression  of  the  organ  during; 
respiration  for  increasing  circulation  in  it ;  at  the  same  time 
promoting  the  secretory  functions  and  metabolic  processes  in 
the  organ,  which  are  very  extensive.  There  is  a  reason  for 
all  the  arrangements  that  obtain  In  short,  the  organ  is  sand- 
wiched, so  to  speak,  between  two  opposing  forces,  notably  the 
diaphragm  and  muscles  in  the  abdomen  ;  while  the  intestines 
serve  as  a  soft,  elastic  cushion  for  effecting  the  rhythmical 
compression  under  the  force  in  the  muscles  during  respiration, 
transmitting  this  upon  it,  and  which,  of  course,  increases  its 
circulation  correspondingly.  And  not  this  only,  but  all  the 
other  actions  as  well,  inclusive  of  the  intestines  and  the 
lymphatics  or  lacteal  system  and  terminal  duct.  It  also  ad- 
mits of  easy  demonstration.  For  this  purpose  a  dog  was  chosen, 
and  several  hours  after  the  animal  had  been  fed  it  was  dis- 
patched, and  the  abdomen  separated  from  the  chest  by  division 
effected  above  the  insertions  of  the  diaphragm  in  the  ribs,  and 
was  completed  by  severing  the  spine  between  the  last  dorsal 
and  first  lumbar  vertebra,  ligating  the  vena  cava,  oesophagus 
and  thoracic  duct  for  preventing  escape  of  the  contents.  As 
will  be  seen,  the  abdomen  separated  from  the  chest  forms  an 
egg-shaped  organ  (Fig.  8b),  the  large  end  of  the  ovoid  looking 
anteriorly,  or  toward  the  chest ;  the  small  ending  posteriorly 
in  the  pelvic  basin  in  which  it  terminates  ;  the  differentiation 
of  this  bony  compartment  having  reference  to  the  crural  sup- 
ports for  sustaining  the  posterior  end  of  the  trunk  during 
locomotion,  and  of  the  body  itself  in  the  erect  position,  as  in. 
man.  The  cava  (B),  oesophagus  (C),  and  aorta,  with  the  accom- 
panying duct  (D),  occupy  the  mid-region  upon  a  line  extend- 
ding  from  the  sternum  to  the  spine,  with  the  cava  a  little  to  the 
right  (left  in  the  picture).  And  everything  being  ready,  the 
stump  of  the  cava  was  then  snipped  with  a  sharp  pair  of' 
scissors.     The  blood  spurted  with  great  force  out  of  the  vessel,, 


222 


THE   FOECE  TO   THE   PORTAL  CIRCULATION. 


leaping  ten  feet,  and  as  though  shot  out  of  a  syringe,  under 
the  force  in  the  abdomen  ;  but  was  not  sustained,  falling  down 
immediately  upon  the  diaphragm  in  a  running  stream.  I  then 
placed  my  open  hand  upon  the  convexity  of  the  diaphragm, 
pushing  it  downward  in  imitation  of  inspiration,  when  the  jet 
was  renewed  instantaneously,  rising  and  falling  with  the 
increase  and  diminution  of  the  pressure  ;  hence,  there  could  be 
no  doubt  on  earth  that  inspiration  rushes  the  venous  blood  in 
the  abdomen  toward  the  lungs.  Of  course,  a  great  deal  of  the 
blood  had  come  from   the  lower  cava  system,  rushing  out  of 


Fig.  83. — Transverse  Section  of  the  Trunk  through  the  Inferior  Margin  of  the  Chest 
around  the  Circle  of  the  Diaphragm,  showing  the  form  of  the  abdomen,  which  is 
egg-shaped,  the  large  end  of  the  ovoid  presenting  in  the  chest-excavation,  the  small 
in  the  pelvic  basin,  which  forms  the  posterior  end  of  the  cavity.  A,  diaphragm  ;  2?, 
stump  of  lower  cava,  ligatured  ;  C,  oesophagus,  ligatured  ;  D,  thoracic  aorta  and 
thoracic  duct,  ligatured  ;  E,  first  lumbar  vertebra  ;  F,  lumbar  muscles  (quadra tus 
lumborum,  etc.). 

both  under  the  action  of  the  elastic  air-cushion,  while  the  force 
transmitted  from  the  containing  walls,  in  imitation  of  the 
action  in  inspiration,  increases  it.  In  the  first  instance,  the 
intra-abdominal  pressure  of  itself  was  sufficient  for  producing 
the  jet,  showing  it  to  be  considerable.  It  should  be  mentioned 
here  that  the  valves  in  the  femoral  veins  obviate  reflux  in  the 
lower  extremities,  so  that  the  systole  in  the  abdomen  produced 
by  inspiration  should  act  as  a  lifting  force  upon  the  blood  in 
the  venous  system,  as  well  as  the  portal  blood,  the  two  being 
affected  simultaneously.     (We  may  remark,  en  passant,  that 


THE    FOKCE  TO   THE  HEPATIC   CIKCl  LATIOJN-.  223 

the  presence  of  valves  in  the  veins  of  the  pelvic  viscera  has  its 
explanation  in  connection  with  the  special  functions  in  these 
organs  for  obviating  reflux  during  expulsive  efforts  which 
will  come  up  further  on.)  In  this  manner,  then,  the  abdo- 
men functions  as  a  great  ventricle  upon  the  venous  blood  for 
forcing  it  toward  the  lungs  during  inspiration.  That  there 
might  be  no  mistake,  however,  about  the  portal  blood  and  the 
blood  in  the  liver,  I  introduced  my  hand  through  an  opening 
in  the  linea  alba  and  compressed,  first  one,  then  another  portion 
of  the  liver ;  and  every  time  I  did  this,  and  the  moment  I  did 
it,  the  blood  spurted  out  of  the  caval  opening,  showing  conclu- 
sively, and  beyond  the  shadow  of  a  doubt,  that  inspiration 
increases  circulation  in  the  liver.  Now,  then,  we  can  .readily 
understand  why  the  hepatic  veins  are  fastened  to  the  liver- 
substance,  being  incorporated  with  it  for  maintaining  patency 
in  the  canals,  as  this  would  favor  venous  efflux  during,  inspi- 
ration, the  blood  rushing  out  of  every  portion  of  the  liver 
directly  into  the  vena  cava,  in  place  of  converging  in  a  common 
trunk,  as  in  the  usual  way ;  the  right,  left  and  central  por- 
tions emptying  their  blood  immediately  into  the  venous  sys- 
tem.    It  is  very  pretty. 

Concerning  the  Relations  which  the  Portal  Vessels  Sustain 
to    the    Hepatic    Veins  for  Producing  Correspondence 
and   an    Uninterrupted   Flow   of    Blood   through   the 
Liver. 
Briefly,  the  portal  vein  is  intercalated  between  two  exten- 
sive capillary  systems — one  in  the  mucous  membrane  of  the 
intestines,  the  other  in  the  liver-substance  ;  consequently,  the 
force  in  the  arterial  system  can  have  but  little  effect  upon  cir- 
culation in  the  liver,  which  must  have  other  force  applied  to 
it  for  making  it  commensurate  with  the  physiological  require- 
ments, and  for  maintaining  correspondence  with  the  absorptive 
processes  in  the  intestines  ;  otherwise  impossible.     This,  as  we 
see,  is  made  dependent  upon  respiration  for  producing  rhyth- 
mical compression  of  the  liver,  and  which  is  supplemented  by 
the  action  in  the  portal  vein  itself,  which  is  very  muscular,  and 
connected  with  respiration  by  means  of  the  rich,  nervous  plex- 
uses embossing  the  vessels  the  same  as  the  arteries,  and,  like 
them  also,  enabling  an  increase  and  decrease  in  the  local 


"22±  THE    FORCE   '10   THE  HEPATIC   CIRCULATION. 

actions  by  expanding  and  contracting  the  lumen,  the  portal 
vein  dividing  up  in  the  liver  exactly  like  an  artery  ;  more- 
over, are  free  to  move  by  reason  of  Glisson's  capsule,  and 
not  fastened  to  the  liver-substance  like  the  hepatic  veins. 
In  short,  they  possess  automatism,  and  being  intimately  con- 
nected with  the  solar  plexus  and  pneumogastric  nerves,  should 
greatly  expedite  circulation  in  the  organ  during  respiration, 
at  the  same  time  it  constitutes  the  fine  adjustment  in  this 
mechanics  in  correspondence  with  the  universal  rule  ;  other- 
wise, the  local  actions  could  not  be  carried  on.  In  other 
words,  the  vessels  have  a  separate  sheath  of  their  own  in 
the  liver,  with  the  nerves  extending  over  the  vessels  the  same 
as  in  the  cavity  of  the  abdomen,  and  which  certainly  jus- 
tifies this  conclusion.  Furthermore,  we  have  seen  that  dur- 
ing inspiration  a  wave  of  contraction  pervades,  first  the 
arterial,  then  the  capillary,  then  the  venous  systems,  which, 
of  course,  would  include  the  portal  vessels,  no  reason  pre- 
senting why  it  should  not  do  so,  but  every  reason  for  it, 
bringing  it  in  correspondence  with  the  systemic  current, 
which  the  scheme  calls  for.  And  since  respiration  is  all-per- 
vading, it  is  hardly  to  be  doubted  that  such  is  the  case.  But 
this  dependence  of  the  portal  circulation  upon  respiration 
calls  for  its  check  and  balance  also,  in  order  to  preserve 
equilibrium,  which  is  furnished  in  the  nervous  plexuses 
that  embrace  the  vessels,  the  strain  to  these,  which  is  occa- 
sioned by  over-distention,  producing  the  reflex  action  for 
relieving  it,  and  known  as  "sighing,"  which  is  simply  an 
effort  to  relieve  an  overcharged  portal  system  ;  in  this  manner 
exciting  the  reflex  actions  in  the  medulla  oblongata,  or  the 
same  as  regulates  the  actions  in  the  lungs  and  in  the  heart,  the 
whole  connecting  through  and  through  ;  otherwise,  correspond- 
ence could  not  be  produced  for  maintaining  a  balance  in  the 
circulation  for  conserving  the  functions.  Its  connection  with 
mental  causes  is  due  to  the  fact  that  profound  mental  disturb- 
ance diverts  nervous  force  from  respiration  for  protracted 
periods,  the  blood  in  consequence  accumulating  in  the  portal 
vessels,  till  finally  comes  the  deep  and  noisy  inspiration  for 
relieving  it,  which  would  also  include  the  venous  stasis  in 
the  systemic  capillaries,  as  this,  too,  is  dependent  upon  res- 


THE    F0ECE  TO   THE  HEPATIC   CIECULATIOlSr.  225 

piration.  But  any  circumstance  that  should  embarrass  res- 
piration, mental  or  mechanical,  would  soon  produce  over-dis- 
tension of  the  portal  vessels,  and  invoke  this  eff ort  for  relieving 
it.  Hence,  full  diet,  with  sedentary  habits,  or  tight  corsage,  as 
in  the  case  of  females,  who  are  notorious  for  these  sounds, 
preventing  free  action  in  the  diaphragm,  consequently  pro- 
ducing engorgement,  while  free  exercise  and  loose  corsage  are 
effectual  remedies.  It  is  very  disagreeable,  the  more  so  be- 
cause deceptive,  since  it  suggests  pain  when  none  exists,  and 
if  the  ladies  should  release  the  abdomen  and  burn  up  the 
internal  bandages,  or  put  them  on  occasionally  when  they  go 
from  home,  taking  more  exercise,  too,  we  would  hear  no  more 
from  them  in  this  direction.  The  whole  trouble  is  in  the 
abdomen,  and  freer  respiration  is  needed  for  relieving  it.  But 
man  has  no  monopoly  of  this  action,  since  the  very  fishes, 
after  remaining  stationary  for  long  periods,  are  seen  to  make 
this  deep  inspiratory  effort,  which  compresses  the  viscera  and 
flares  open  the  opercula  and  branchiae  at  one  and  the  same 
time,  for  relieving  venous  stasis,  the  principle  in  the  mechanics 
being  fundamentally  the  same  as  in  the  higher  animals,  due 
allowance  being  made  for  the  stage  in  development.  The 
connection  of  the  muscles  in  the  abdomen  with  the  lower 
jaw  causes  it  and  the  opercula  to  fly  widely  open  at  one  and 
the  same  time,  when  the  abdomen  contracts  for  increasing 
pressure,  thereby  increasing  the  action  in  the  branchiae,  all 
the  parts  being  fully  coordinated  in  the  medulla  oblongata. 
The  horse  and  ox,  standing  quietly  in  full  digestion,  are  seen 
every  now  and  then  to  take  one  of  these  deep  inspiratory 
efforts,  rocking  forward  so  suddenly  upon  their  feet  as  to 
rouse  them  from  the  doze.  Anything  which  should  hasten 
respiration,  therefore,  such  as  singing,  laughing,  talking  or 
bodily  exercise,  would  greatly  expedite  the  portal  circulation. 
Hence  the  trite  saying  ' '  Laugh  and  grow  fat "  is  a  truism 
in  physiology,  since  the  effect  of  the  former  is  to  pump  the 
portal  blood  and  lymph  into  the  systemic  current  and  produce 
fullness  in  the  vessels,  which  favors  the  nutritive  processes  and 
the  formation  of  fat.  It  was  a  sharp,  true  saying.  The  effect 
of  the  rapid  pumping  action  in  the  abdomen  in  reducing  the 
volume  of  the  portal  blood  has  forcible  illustration  in  the 


226 


PHYSIOLOGICAL    ANATOMY 


vocalist,  who  complains  of  "•  a  feeling  of  emptiness  in  the 
stomach,"  and  rushes  to  the  restaurant  to  till  himself  full 
again,  eating  ravenously  to  appease  the  appetite.  The  deep 
and  long  inspiratory  efforts,  followed  by  the  long  forced  expi- 
rations, had  literally  emptied  the  portal  circulation,  as  it  were, 
also  the  viscera  of  the  force-producing  elements.  In  fine,  for 
increasing  the  portal  circulation  it  calls  for  rhythmical  com- 
pression of  the  liver  and  portal  vessels  synchronous  with  respi- 
ration. 

The  following  beautiful  cuts  (Figs.  84,  85)  show  the  mode  of 
termination  of  the  portal  vessels  in  the  inter-lobular  veins,  and 


Fig.  84. — Rabbit's  Liver  Injected,  showing  a  portal  branch,  the  vence  inter-lobitlares,  the 
capillary  network,  and  a  vena  intra-lobularis  in  the  centre  of  a  lobule. — Frey. 

these  again  in  the  intra-lobular  (Fig.  85,  3,  2,  1),  or  radicals  of 
the  open  hepatic  veins  ;  so  that  one.  can  readily  understand 
why  rhythmical  compression  of  the  liver  should  increase  cir- 
culation in  the  venous  channels,  while  fhe  speed  of  the  cur- 
rent thus  produced  would  depend  upon  the  energy  in  the 
respiratory  rhythms.  Of  course,  the  action  in  the  portal 
vessels  should  be  considered  as  in  connection  with  this, 
while  the  increase  of  pressure  in  the  parenchymatous  tissue 
promotes  absorption,  but  increase  in  arterial  pressure  exerts 
no  influence  upon  it  (■  enersich)  ;*  hence,  rhythmical  com- 

*  Ludwig's  Arbeiten,  1871,  p.  53. 


PH    SIOLOGICAL    A1STAT0MY.  227 

pression  of  the  liver  during  respiration  is  essential  for  expedit- 
ing absorption  in  the  liver  for  maintaining  this  in  corre- 
spondence with  the  absorptive  processes  in  the  intestines ; 
otherwise  impossible.  In  this  manner,  then,  the  portal  circu- 
lation is  maintained  in  correspondence  with  the  absorptive 
processes  in  the  intestines  and  liver,  the  pressure  produced  by 
the  gases,  together  with  the  respiratory  rhythms,  acting  upon 
both  alike  for  compelling  this  circumstance.  In  other  words, 
the  fluids  in  the  parenchyma  are  pumped  into  their  respective 
channels  simultaneously  with  the  fluids  in    the  intestines. 


Pig.  85. — Transverse  Section  of  a  Single  Hepatic  Lobule.  — Sappey.  1 ,  Intra-lobular  vein, 
cut  across  ;  2,  2,  2,  2,  afferent  branches  of  the  intra-lobular  vein  ;  3,  3,  3,  3,  3,  3,  3, 
3,  3,  inter-lobular  branches  of  the  portal  vein,  with  its  capillary  branches,  forming 
the  lobular  plexus,  extending  to  the  radicles  of  the  intra-lobular  vein. 

Finally,  to  this  mechanics  must  be  added  the  action  in  the 
heart,  the  diastoles  in  the  right  side  serving  to  aspirate  the 
hepatic  veins,  at  one  and  the  same  time  that  they  aspirate  the 
venous  system,  the  vessels  also  debouching  in  the  cava  close  to 
the  heart,  whereby  an  uninterrupted  flow  is  maintained  in  the 
organ,  the  same  as  in  the  lungs,  but  is  not  sufficient  to  make  it 
uniform,  the  blood  tending  to  accumulate  during  expiration  ; 
and  which  affords  opportunity  for  effecting  metamorphosis, 
an  amount  of  time  being  required  for  this  purpose,  which  is 
also  in  correspondence  with  the  action  in  the  lungs — indeed, 
in  every  organ. 


228  MUSCULAR  FORCE   FOR  THE  BILE. 

Concerning  the  Action  in  Bile. — Of  course,  the  same  mechan- 
ics apply  for  compelling  the  bile  through  the  gall  ducts  as  the 
blood  through  the  hepatic  veins,  the  force  in  respiration  affect- 
ing both  alike  and  simultaneously.  But  here  also  a  special 
force  applies  for  increasing  the  actions,  ard  interesting  ana- 
tomical dispositions  obtain  for  separating  the  bile  fiom  the 
blood,  which  it  would  be  as  well  to  mention  in  this  connection. 
According  to  Professor  Hering,  the  gall  capillaries  or  intra- 
lobular gall  ducts  do  not  run  along  the  boundary  walls  of  the 
liver  cells,  but  are  incorporated  with  them,  blending  with  the 
walls  in  which  they  lose  themselves,  as  it  were  ;  thus  bringing 
them  into  intimate  relations  with  the  cell  contents.  Further- 
more, that  the  blood  capillaries  occupy  the  spaces  between  the 
cells  upon  the  opposite  side,  the  cell-substance  intervening 
between  them,  the  effect  of  the  arrangement  being  to  isolate 
the  gall  capillaries,  separating  them  completely  from  the  blood 
capillaries  ;  so  that,  when  pressure  is  made  upon  the  liver  dur- 
ing inspiration,  the  blood  passes  in  one  direction,  the  bile  in 
quite  another,  or  opposite  direction.     It  is  very  pretty. 

Now,  then,  to  this  must  be  added  the  special  adjustments  in 
the  ducts  for  compelling  movement  in  the  bile,  which  in- 
volves automatic  action  in  this  tubular  system,  the  same  as 
the  portal  vessels  and  hepatic  artery,  with  which  they  are 
inclosed  inGrlisson's  capsule,  since  they  also  require  independ- 
ent movement  in  connection  with  their  special  functions,  or 
the  power  to  expand  and  contract  upon  the  contents,  which, 
of  course,  is  intiuenced  by  nervous  force  propagated  from  the 
intestinal  mucous  surface  during  digestion,  for  compelling  the 
secretions  into  the  duodenum,  at  once  the  purpose  of  this 
arrangement.  But  as  the  biliary  secretion  is  going  on  all  the 
while,  we  have  a  special  arrangement  in  the  gall-bladder  for 
storing  the  bile  in  the  interim  of  digestion  ;  and  this  mechan- 
ics also  contains  several  very  interesting  features.  Thus,  the 
gall-bladder  occupies  an  elevated  position  against  the  liver- 
substance,  to  which  it  is  fastened  by  an  overlying  layer 
of  peritoneum  and  connective  tissue  fibres,  while  its  neck 
is  occupied  by  a  spiral  valve  (Fig.  8  ),  the  whole  whipped 
off  to  one  side  of  the  main  hepatic  duct.  Hence,  when 
the  orifice  of   the  duct  closes   during   the  interim   of  diges- 


MUSCULAR  FORCE  FOR  THE  BILE. 


229 


tion  the  effect  would  be  to  cause  reflux  of  the  bile  up  the 
cystic  duct,  the  force  in  the  diaphragm  compelling  this 
circumstance.  Muscles  and  nerves  to  the  gall-bladder  also. 
Why  %  As  we  have  already  said,  muscles  and  nerves  are  the 
provision  for  producing  force ;   hence,  they  must  relate  to  the 


Fig.  86. — Longitudinal  Section  of  Duodenum,  showing  tne  crescentic  folds  of  the 
valvulae  conniventes,  pancreatic,  hepatic  and  cystic  ducts,  together  with  the  spiral 
valve  at  the  neck  of  the  gall-bladder. — Gray. 

bile,  for  compelling  it  into  the  cavity  during  the  interim  of 
digestion,  and  for  compelling  it  into  the  intestines  in  corre- 
spondence with  the  functions  in  the  latter,  muscular  and 
nervous  force  being  applied  all  along  the  line  for  increasing 
the  actions  and  producing  correspondence  ;  otherwise,  this  also 


230  AUTOMATISM    IN    THE   G ALL-DUCTS. 

would  be  meaningless.  According  to  Iltnle,  the  coats  of  the 
gall-bladder  are  formed  of  layers  of  connected  tissue,  alternating 
with  lamina?  of  unstriped  muscles,  the  fibres  crossing  each  other 
in  all  directions,  the  nerves  for  the  most  part  springing  from 
the  solar  plexus.  It  results  from  this  arrangement  in  the 
parts  that  the  gall-bladder,  inclusive  of  the  ducts,  must  be  in- 
fluenced by  resrjiration  and  the  local  actions,  being  under  con- 
trol of  the  central  nervous  system  and  the  local  ganglia.  And 
by  looking  closely  into  the  mechanics,  it  is  also  easy  to  per- 
ceive how  the  action  in  the  diaphragm  should  materially  assist 
the  ascent  of  the  bile  up  the  duct  into  the  bladder.  Notably, 
the  gall-bladder  is  fastened  to  the  liver  while  the  terminal  end 
of  the  common  duct  is  inclosed  in  the  walls  of  the  duodenum, 
which  is  firmly  secured  to  the  spine  by  connective  tissue,  and 
the  overlying  peritoneum  (for  it  would  not  do  to  have  it  sag- 
ging by  the  ducts) ;  hence,  the  to-and-fro  movements  in  the 
diaphragm  with  the  mouth  of  the  duct  closed  must  produce 
reflux  of  the  bile  into  the  gall-bladder,  the  spiral  valve  serv- 
ing to  assist  the  action,  at  the  same  time  that  it  functions  as 
the  floor  of  support.  It  follows,  however,  that  the  ducts  must 
contract  and  elongate  with  inspiration  and  expiration  in  order 
to  obviate  strain,  otherwise  inevitable,  since  the  diaphragm 
oscillates  as  much  as  two  inches,  and  which  could  not  other- 
wise than  affect  the  movement  of  the  bile.  At  the  same  time, 
we  must  remember  that  there  is  automatic  action  in  connec- 
tion with  the  flow,  since  the  functions  in  the  gut  call  for  this, 
the  sensory  impressions  produced  in  the  mucous  membrane 
by  the  food  being  transmitted  to  the  special  local  ganglia  for 
the  purpose.  It  is  all  connected  by  nervous  force  for  effecting 
coordination  and  producing  correspondence  in  the  local  actions. 
The  bile,  then,  flows  down  the  ducts  into  the  gut  during 
digestion,  responsive  to  the  calls  made  upon  it ;  while  during 
the  interim  it  flows  up  the  cystic  duct,  while  the  bladder  ex- 
pands pari  passu  with  the  afflux,  the  same  as  the  urinary 
bladder,  or  from  sensory  impressions  in  the  mucous  membrane 
produced  by  the  bile.  Hence,  it  is  easy  to  perceive  how  it 
should  become  enormously  distended  with  the  bile  when  greatly 
in  excess  of  the  wants  in  the  digestive  processes ;  not  that  it 
is  forcibly  distended  by  the  action  in  the  diaphragm  and  gall- 


RHYTHMICAL   COMPRESSION   OF   THE  PANCREAS.  231 

ducts,  any  more  than  the  urinary  bladder  is  distended  by  the 
action  in  the  ureters  and  kidneys,  which  cannot  be  thought 
of  ;  but  that  the  organ  expands  till  it  can  expand  no  more, 
and  pain  is  produced  by  the  traction  upon  the  nervous  fila- 
ments. We  might  mention  here  the  existence  of  the  close 
capillary  plexuses  in  the  mucous  membrane  of  the  gall-blad- 
der, which  would  effect  rapid  absorption,  so  that  the  bile  soon 
becomes  thick  and  viscid  ;  hence,  cannot  be  retained  long  at  a 
time  without  danger  of  becoming  inspissated,  producing  gall- 
stones. 

Concerning  the  Action  in  the  Pancreas. — Taking  up  the 
circumstances  in  anatomy  as  we  go  along,  the  pancreas  comes 
next  in  order,  its  duct  emptying  into  the  bile  duct  (sometimes 
opening  independently  in  the  gut),  the  separate  secretions  thus 
intermingling  in  the  common  duct  previous  to  debouching  in 
the  duodenum  (Fig.  86).  Why  the  two  fluids  should  be  thus 
intimately  admixed,  bringing  them  together  as  speedily  as 
possible  before  undergoing  admixture  with  the  chyme,  must 
have  reference  to  interchanges  between  them,  whereby  they 
gain  in  effectiveness,  which  would  be  the  natural  explanation  ; 
but  what  they  are,  chemical  or  otherwise,  1  am  not  prepared 
to  say,  only  throwing  out  the  suggestion  that  the  circumstance 
gives  rise  to.  The  organ  is  conical  in  shape,  the  broad  end  or 
base,  known  as  the  "head,"  abutting  against  the  gut,  which 
bends  around  it,  above  and  below,  embracing  it  (Fig.  b7),  while 
the  long,  tapering  body  extends  across  the  crura  of  the  dia- 
phragm behind  the  stomach  to  the  spleen  attached  to  the 
great  end  or  cut  de  sac  of  the  stomach,  where  it  terminates  in 
the  tail.  It  results  from  this  arrangement  in  the  parts  that 
the  organ  is  rhythmically  compressed  during  respiration,  the 
same  as  the  liver,  both  simultaneously,  which  should  cause 
the  secretions  to  flow  out  at  one  and  the  same  time  with  the 
bile,  and,  like  it,  also  flowing  from  high  to  low  pressure,  since 
the  churning  action  in  the  duodenum  connected  with  its  func- 
tions must  inevitably  determine  low  pressure  around  the  ter- 
minal duct  with  every  expansion  in  the  gut.  Then,  again,  the 
churning  action  in  the  stomach  itself  should  effect  changes  of 
pressure  in  the  gland  for  promoting  the  discharge  of  the  secre- 
tions. 


232 


AUTOMATISM  IN  THE  PANCREAS. 


Finally,  we  have  to  mention  the  automatism  in  the 
gland  cells  and  discharging  ducts,  without  which  nothing 
could  be  accomplished  in  the  secretory  processes  ;  while  this 
in  turn  is  connected  with  the  action  in  the  gut  by  means  of 
the  special  nerves  and  ganglia  for  unifying  and  coordinating 
the  movements,  the  same  as  the  liver  and  bile  ducts,  whereby 
uniformity  is  produced  throughout,  otherwise  impossible  ;  the 
pancreatic  secretions  being  controlled  by  sensory  impressions 
in  the  mucous  membrane,  produced  by  the  food,  the  same  as 


Fig.  87.— Pancreas  and  its  Relations.— Gray. 


the  hepatic,  and  salivary  glands  with  which  it  is  homologous, 
the  structure  being  the  same  or  racemose.  All  of  which  is 
plain  enough. 

Concerning  the  Action  in  the  Spleen. — That  the  spleen  pos- 
sesses certain  powers  of  expanding  and  contracting  in  connec- 
tion with  the  functions  in  the  organ,  there  is  small  room  for 
doubt,  for  the  following  reasons — namely:  1.  The  investing 
capsule  forming  the  framework  of  the  oraan  (giving  off  the 
cords  or  trabecular,  in  which  the  parenchyma  or  splenic  pulp  is 
situated — being  traversed  and  inclosed  by  the  trabecular — and 


PHYSIOLOGICAL   ANATOMY   OF  THE   SPLEEN.  233 

passing  in  at  the  hilus  with  the  vein  and  artery,  incloses  them 
also)  is  homologous  with  arterial  structure,  being  composed 
of  fibro-elastic  tissue  and  muscles.  Elastic  tissue  fibres  are 
abundantly  intermixed  with  connective  tissue  fibres  in  the 
outer  or  external  portions  of  the  capsule,  while  the  deeper 
or  internal  portions  are  rich  in  unstriped  muscle  fibres, 
which  are  interspersed  with  the  cords  or  trabecule  through  the 
parenchyma,  and  being  inserted  into  the  walls  of  the  veins 
with  which  the  trabeculse  are  incorporated,  it  is  manifest  that 
contraction  of  those  fibres  would  have  the  effect  of  flaring  open 
the  veins,  and  expansion  the  oppposite  effect ;  while  contrac- 
tion of  the  envelope  itself  should  condense  the  spleen  and 
drive  out  the  blood  in  the  veins,  expansion  would  have 
the  opposite  effect.  But  with  the  actions  coordinated  would 
make  it  still  more  effective  for  compelling  the  venous  blood  in 
and  out  of  the  organ;  and  the  terminal  ends  of  the  veins 
being  open,  the  whole  cell- structure,  the  lymphoid  follicles, 
and  Malpighian  corpuscles  are  thus  bathed  in  the  portal  blood 
brought  into  them  by  this  action  in  the  gland.  In  short,  there 
must  be  some  reason  for  the  presence  of  elastic-tissue  fibres 
and  muscles  in  the  gland,  and  the  special  relations  they  sustain 
to  the  veins. 

2.  The  walls  of  the  veins,  where  they  enter  the  hilus,  and  for 
some  distance  in,  are  very  thick  and  dense,  by  reason  of  being 
incorporated  with  the  trabeculse,  blending  together ;  while  in 
the  case  of  the  artery  and  its  branches,  the  walls  are  not  joined 
to  the  trabeculse,  but  the  vessels  are  contained  in  a  loose 
sheath  of  areolar  tissue,  thus  allowing  free  action  in  them, ' 
which  is  essential  to  the  life  in  the  cell-broods  of  the  lymphoid 
follicles  and  Malpighian  corpuscles,  enabling  them  to  increase 
or  diminish  the  supply  of  arterial  blood  for  maintaining  the 
life  that  is  in  them,  and  fulfilling  their  proper  functions  ;  in- 
creasing or  diminishing  the  actions  upon  occasion  in  response 
to  nervous  stimulus,  in  correspondence  with  the  universal 
method.  In  short,  there  must  be  some  special  reason  for  the 
anatomical  dispositions  that  obtain  in  the  arteries  as  well  as  in 
the  veins,  for  they  cannot  be  purposeless. 

3.  The  great  relative  size  of  the  veins,  which  is  greatly  in 
excess  of  the  arteries,  the  same  applying  for  the  trunks  of  the 


234  PHYSIOLOGICAL   ANATOMY    OF   THE   SPLEEN. 

vessels,  the  splenic  vein  being  a  number  of  times  larger  than 
the  artery  ;  there  must  be  some  reason  for  this  also,  for  it 
neither  can  be  purposeless.  And  there  can  be  but  little  doubt 
that  the  organ  functions  as  a  reservoir  for  relieving  an  over- 
distended  portal  system,  expanding  during  this  time  under 
special  stimulus,  and  thus  diverting  the  blood  from  the  liver, 
and  so  relieving  it.  It  would  account  for  the  elastic  tissue  fibres 
and  the  muscles  in  the  spleen,  together  with  the  special  arrange- 
ments in  the  walls  of  the  veins  in  the  spleen  Furthermore, 
we  know  the  organ  does  rayjidly  increase  in  size  in  acute 
febrile  disorders,  from  the  congestions  they  produce  in  the 


Fig.  88. — Transverse  Section  of  the  Human  Spleen,  showing  the  distribution  of  the 
splenic  artery  and  its  branches. — Gray. 

liver  and  portal  system ;  nor  is  it  possible  for  this  condi- 
tion to  be  produced  by  the  action  in  the  vein  simply,  the 
organ  absolutely  inhibiting  it ;  the  walls  of  the  vein  yield- 
ing long  before  the  dense  structures  in  the  spleen.  Besides, 
it  would  involve  the  whole  portal  system.  It  cannot  be 
thought  of.  The  sharp  pain  in  the  left  side,  induced  by 
running,  has  its  explanation  in  an  over-distended  spleen,  the 
organ  rapidly  expanding  for  relieving  a  surcharged  liver  and 
portal  system,  thereby  pulling  upon  the  nervous  filaments  and 
producing  pain,  just  as  an  over-distended  bladder  pioduces 
pain,  only  the  one  is  more  rapidly  produced  than  the  other, 
and  the  pain  sharper.      Finally,  the  simple  experiment  of 


MUSCULAR    FOECE    FOE  THE   SPLEEN. 


235 


dipping  the  spleen  into  warm  water  condenses  the  organ  ;  the 
galvanic  current  also  ;  though  by  reason  of  the  density  of  the 
structure  the  action  is  necessarily  slight ;  hence,  there  can  be 
no  doubt  that  it  also  expands.  Moreover,  the  vessels  are  em- 
bossed by  a  dense  plexus  of  nerves  connecting  with  the  solar 
plexus,  which  at  once  shows  that  there  must  be  great  activity 
in  the  organ  ;  otherwise,  it  were  all  meaningless.  The  special 
anatomy  is  deeply  interesting,  but  it  is  difficult  to  produce 


Fig.  89.— Oae  of  the  Splenic  Corpuscles,  showing  its  relations  with  the  blood  vessels. — 
G-ray.  a,  Arterial  twig,  embracing  a  Malpighian  corpuscle.  The  terminal  branches 
of  the  splenic  vein  are  seen  losing  themselves  in  the  corpuscle,  in  which  they  open 

a  mental  picture  of  it ;  indeed,  altogether  impossible,  though 
some  general  idea  may  be  formed.  The  lymph  follicles,  cells 
and  Malpighian  corpuscles  are  in  close  and  intimate  relations 
with  the  arterial  vessels,  as  a  matter  of  course,  the  arterial 
capillaries  surrounding  them  (Fig.  88).  The  manner  in  which 
the  veins  lose  themselves  in  the  Malpighian  corpuscles,  and  the 
intimate  relations  they  sustain  to  the  arterial  feeder,  are  very 
forcibly  shown  in    the  following  beautiful  cut  (Fig.  89,  a). 


236  ABSENCE   OF  VALVES   IN   THE   PORTAL   SYSTEM. 

For  the  rest,  the  reader  is  referred  to  the  literature  of  the  sub- 
ject and  works  on  histology. 

The  portal  system  is  brought  into  prominence  in  the  follow- 
ing beautiful  cut  (Fig.  (J0)  for  impressing  the  special  anatomy 
upon  the  mind.  It  would  seem  a  bold  thing  to  disconnect 
the  viscera  from  the  containing  walls  in  order  to  increase  the 
action  in  them,  and  to  send  the  vessels  and  nerves  to  the 
organs  over  floating  ligaments  ;  but  a  bolder  one  to  break  up 
the  portal  veins  into  a  second  capillary  system  in  the  liver, 
threading  it  in  every  direction  and  supplying  every  cell  with 
the  fluid,  and  as  though  the  vein  were  itself  an  artery  with  the 
force  of  the  heart  at  its  root,  as  in  the  aorta,  whereas  the 
heart  is  far  removed,  the  vessels  floating  in  a  ligament  and  a 
capillary  barrier  between,  to  be  re-collected  through  the  radi- 
cals of  the  hepatic  veins  that  debouch  in  the  lower  cava,  and 
to  send  the  vast  bulk  of  the  alimental  fluids  over  such  a  route, 
the  organism  itself  depending  upon  it :  there  is  a  problem  for 
you,  but  all  worked  out  and  open  for  inspection.  But  the 
solution  is  only  possible  upon  the  basis  of  the  fundamental 
law  underlying  the  organism  and  autonomy  in  the  structures, 
the  local  actions  intimately  blending  with  the  systemic 
mechanics,  by  means  of  the  nerves  leading  out  of  the  solar 
plexus,  in  which  the  visceral  nerves  converge  (Fig.  108,  5,  2,  1 ), 
this  connecting  with  the  respiratory  and  vaso-motor  centres 
through  the  pneumogastric  (Fig.  109,  2)  and  great  splanchnic 
nerves  (3,  3),  together  with  the  pumping  actions  in  the  lungs, 
heart,  arteries  and  capillaries,  inclusive  of  the  force  in  the  ven- 
ous system  ;  the  whole  coordinated  through  the  medulla  oblon- 
gata, whirls  it  over  the  road-bed  in  the  vascular  lines  with  such 
expedition  as  to  make  the  journey  within  a  minute  from  the 
cavity  of  the  stomach.  But  all  plain  enough  in  the  light  of  the 
law  that  applies  and  automatism  in  the  tissues,  the  muscular 
and  nervous  forces  swelling  at  the  difficult  points  for  compelling 
the  passage  and  completing  the  circuit ;  otherwise  is  utterly 
inexplicable. 

There  is  more  !  In  all  the  expanse  of  venous  territory  in 
the  portal  system,  not  a  valve  presents.  Why  (  The  answer 
is  not  far  to  seek.  The  presence  of  a  valve  in  the  portal  vein 
would  prevent  reflux,  so  that  congestion  of  the  liver  could 


ABSENCE    OF   VALVES    IN    THE   POKTaL    SYSTEM. 


237 


not  find  relief  in  copious  transudations  in  the  intestiual  canal, 
producing  diarrhoea,  thereby  conserving  liver-structure,  which, 
is  the  object  of  this  arrangement,  and  would  inevitably 
end  in  disaster.  Indeed,  we  have  example  of  this  in  the 
partial   constrictions   which  occur  in  the  liver  from  hyper- 


Fig.  90. — Portal  System  in  Diagram. — Gray.  The  transverse  colon  is  removed,  the- 
duodenum  divided  near  the  pyloric  orifice  and  the  stomach  pulled  aside,  in  order  to 
expose  the  vena  portse,  with  the  mesenteric  arteries  omitted,  for  better  definition  of 
the  mesenteric  veins. 

plasia  of  connective  tissue  produced  by  alcohol,  interfering 
with  the  course  of  the  blood  through  the  liver,  tending  to 
dam  it  back  in  the  portal  system,  leading  to  diarrhoea  and. 
effusions  in  the  cavity  of  the  abdomen,  producing  the  drunk- 
ard's dropsy.  But  with  valves  in  the  veins,  this  reflux  could- 
not  occur,  and  life  would  have  a  speedy  termination,   clos- 


238  ABSENCE   OF   VALVES   IN   THE   PORTA  L   SYSTEM. 

ing  upon  it,  sharp  and  terrible  as  the  jaws  of  a  mastodon.  It 
is  well,  then,  for  the  drunkard  that  there  are  not  valves  in  the 
portal  vein;  while  temporary  congestions  of  the  liver  are 
passed  over  and  never  known,  finding  prompt  relief  in  copious 
evacuations  through  the  bowels.  And  it  is  no  wonder  that 
purgatives  are  so  much  lauded,  since  overfeeding  is  extremely- 
liable  to  overload  the  portal  system,  while  purgatives  effect 
rapid  depletion,  thereby  restoring  the  balance,  and  everything 
runs  on  as  before.  JSrot  at  all  strange.  But  it  would  be  far 
better  to  stop  short  of  gluttony,  and  not  overrun  the  capacity 
in  the  mechanics.  Abernethy  was  very  close  to  the  mark : 
"Half  of  the  diseases  come  from  fretting,  the  other  half  from 
stuffing."'  The  fir.-t  we  can  readily  understand,  from  the 
effects  it  must  inevitably  exert  upon  the  local  and  systemic 
actions,  from  the  diversion  of  nervous  force  it  must  inevitably 
produce,  tending  to  engorgements  and  disaster ;  the  second 
has  already  been  considered. 

In  conclusion :  The  soft,  compressible  portal  veins  spread 
out  in  the  viscera  could  not  otherwise  than  undergo  rhyth- 
mical compression  during  respiration,  which  should  tend  to 
force  the  blood  along  the  channels  in  the  liver  with  corre- 
sponding energy ;  this,  together  with  the  actions  in  the  ves- 
sels in  connection  with  respiration  and  the  functions  in  the 
liver,  constitutes  a  dual  foive  for  speeding  the  blood  through 
the  liver,  comprising  the  coarse  and  fine  adjustments  in  this 
mechanics  in  correspondence  with  the  universal  rule,  muscu- 
lar and  nervous  force  applying  everywhere  for  speeding  it 
through  the  channels.  And  with  this  arrangement  in  the 
parts,  we  can  readily  understand  the  rapidity  in  the  absorptive 
processes  in  the  stomach  ;*  otherwise  is  utterly  inexplicable. 
The  force  in  the  stomach  for  compelling  it  in  the  vessels,  the 
force  in  the  walls  of  the  abdomen  and  portal  vein  for  com- 
pelling it  through  the  liver,  and,  finally,  the  force  in  the  pump- 

*  Thus,  in  a  case  of  extra  version  of  the  bladder  observed  by  Professor  Erich- 
sen  (Medical  Gazette,  Vol.  XXVI.,  p.  363),  in  which  the  urine  could  be  col- 
lected immediately,  it  was  found  that  when  a  solution  of  ferrocyanide  of 
potassium  was  taken  into  the  stomach,  the  salt  was  detected  in  the  urine  in 
one  instance  within  one  minute,  and  in  three  other  instances  within  two  and 
a  half  minutes. 


.A<TI0N   in  the  gullet  during  respiration. 


239 


ing  actions  in  the  lungs,  heart  and  arteries,  inclusive  of  the 
venous  system  for  completing  the  circuit — it  is  all  plain 
enough  and  easily  understood.  But  in  the  absence  of  this  law, 
night  comes  again,  and  utter  darkness  pervades  it  all. 

Adjustments  Necessitated  by  the  Action  in  the  Diaphragm* 
- — We  have  now  to  mention  a  number  of  adjustments  in  the 
organs  necessitated  by  the  action  in  the  diaphragm,  notably 
the  portal  vein  lower  cava,  diaphragm,  and  oesophagus.   Begin- 


Fig.  91.— Lateral  Section  of  the  Chest  and  Removal  of  Left  Lung,  showing  the  point  at 
which  the  gullet  pierces  the  diaphragm-  A,  diaphragm  ;  C,  oesophagus  ;  B,  heart, 
showing  through  the  pericardium,  with  the  phrenic  nerve  (left)  running  over  it  to 
reach  the  diaphragm  ;  D,  arch  of  aorta  ;  E,  left  bronchus  ;  F,  left  phrenic  nerve  ; 
G,  superior  vena  cava  ;  H,  left  pulmonary  artery. 

ning  with  the  last,  attention  is  directed  to  the  manner  in  which 
the  oesophagus  penetrates  the  diaphragm,  which  is  through 
the  most  active  portions,  entering  it  at  its  greatest  convexity 
(Fig.  91,  A,  C),  and  expanding  at  once  to  form  the  stomach  ; 
the  cardiac  orifice  fitting  close  up  against  the  diaphragmatic 
opening,  while  the  portion  included  in  the  diaphragm  is  the 
narrowest  portion  of  the  gullet ;  furthermore,  is  secured  to  the 
diaphragm  by  means  of  connective  tissue  fibres  and  the  close- 
fitting  muscular  collar  in  the  diaphragm  (Fig.  92,  iV"),  so  that 
there  can  be  no  slipping  up  and  down  of  the  collar  upon  the  gul- 
let during  respiration.    It  results,  therefore,  that  the  oesophagus 


240  THE  ACTION    IN"   THE   MUSCULAR   COLLAR. 

elongates  during  inspiration  and  contracts  during  expiration, 
while  coordination  is  effected  by  means  of  the  pneumogastric 
nerves,  as  may  be  readily  inferred,  being  the  nerves  of  motion 
to  the  organ.  The  great  number  of  the  longitudinal  fibres 
has  its  explanation  in  this  circumstance ;  at  the  same  time 
are  of  service  in  passing  the  bolus,  the  expansions  and  con- 
tractions in  deglutition  involving  elongation  and  contraction  of 
the  tubing.  Magendie  had  noticed  that  the  lower  third  of  the 
oesophagus  remained  firmly  contracted  for  about  thirty  sec- 
onds after  the  bolus  had  passed  into  the  stomach,  feeling- 
like  a  cord  firmly  stretched ;  the  object  of  this,  of  course, 
being  to  afford  the  requisite  time  for  equalizing  pressure  in  the 
stomach  by  means  of  the  reflex  actions  propagated  through  the 
medulla  oblongata  to  the  muscles  in  the  abdomen,  as  before 
remarked  ;  and  the  action  continues  during  digestion  for  the 
reason  that  the  churning  action  in  the  stomach  tends  to  regurgi- 
tation. This  is  proven  by  the  fact  that  the  contract  ion  is  always 
increased  by  pressing  the  stomach  with  the  object  of  forcing 
some  of  the  contents  through  the  cardiac  orifice.  JSTow,  then, 
since  inspiration  increases  pressure  in  the  stomach  and 
abdomen,  we  can  readily  understand  the  significance  of  the 
muscular  collar  in  the  diaphragm,  and  why  constriction  of  the 
oesophagus  should  correspond  with  the  depth  of  inspiration, 
the  one  calling  for  the  other  to  obviate  regurgitation.  Further- 
more, why  there  should  be  suspension  of  respiration  during 
deglutition,  in  order  to  effect  relaxation  or  expansion  in  the 
muscles  of  the  diaphragm,  which  not  only  effects  reduction 
of  pressure  upon  the  gullet,  but  in  the  stomach  and  abdomen 
as  well.  Thus,  all  this  mechanics  is  easily  understood.  The 
several  openings  in  the  diaphragm  involve  a  number  of  very 
pretty  adjustments,  well  worthy  of  undivided  attention — are 
simply  wonderful.  Thus,  taking  the  powerful  muscular  bun- 
dles that  embrace  the  oesophagus,  and  looking  from  their 
points  of  origin  in  the  tendinous  centre  of  the  diaphragm  to 
their  points  of  insertion  in  the  lumbar  vertebrae,  it  will  be  seen 
that  they  form  two  arcs  around  the  oesophagus  and  aorta  (Fig. 
92),  much  in  the  form  of  the  figure  8,  so  as  to  form  the  ellip 
tical  openings,  or  the  oesophageal  and  aortal  respectively. 
Now,  then,   since  the  muscles  contract  upon  a  line  between 


THE   ACTIO N-   I]ST   THE   PILLAES    OF   TIIK   DIAPHRAGM. 


2ll 


their  origin  and  insertion,  both  openings  would  be  occluded 
during  inspiration  but  for  the  following  disposition  of  the 
fibres ;  notably  :  1st.  The  arc  upon  the  left  side  of  the  aorta  is 
bounded  by  a  tendon,  which  is  the  point  of  insertion  for  the 


t^flfom  ,  Cartilo-t 


•      for  tenser 


S/Janeb'tc 


Pig.  92.— The  Diaphragm  as  Viewed  from  Below.— Oray.     The  letters  A,  B,  C,  D,  E, 

N,  O,  are  added. 

thick  layers  of  muscular  fibres  proceeding  from  the  left  leaflet 
of  the  central  tendon  of  the  diaphragm  (A,  B),  so  that  during 
inspiration  this  is  pulled  away  from  the  aorta  at  the  same  time 
that  it  compresses  the  oesophagus  upon  the  right  in  the  upper 
arc.  2d.  The  arc  upon  the  right  of  the  aorta  is  also  bounded 
by  a  tendon,  which  is  the  point  of  insertion  for  the  thick  mus- 


242  PHYSIOLOGICAL   ANA'IOMT   IX   THE   DIAPHEAGM. 

cular  layers  proceeding  from  the  right  leaflet  of  the  central 
tendon  (C,  i>),  so  that  during  inspiration  this  is  pulled  away 
from  the  aorta  at  the  same  time  that  it  compresses  the 
oesophagus  upon  the  left  in  the  upper  arc.  It  results  from  this 
arrangement,  therefore,  that  the  aorta  is  relieved  from  pressure 
during  inspiration,  while  there  is  progressive  increase  of  press- 
ure upon  the  oesophagus,  the  necessity  for  which  has  already 
been  given. 

Finally,  looking  from  the  aortic  to  the  vena  caval  opening, 
we  find  that  the  same  principle  in  mechanics  is  maintained  here 
also.  For  example,  this  opening  occurs  at  the  point  of  junc- 
ture of  the  middle  and  right  leaflets  in  the  central  tendon, 
while  the  muscular  layers  that  have  insertion  in  the  right  aor- 
tic arc  have  their  points  of  origin  upon  the  left  of  the  vena  cava 
opening  (Fig.  92) ;  hence,  when  these  muscles  contract  during 
inspiration,  they  must  inevitably  pull  upon  the  left  side  of  this 
1  nin en  and  flare  it  open,  at  the  same  time  relieving  the  aorta 
by  pulling  upon  the  lower  arc.  Then,  again,  upon  the  right 
or  external  side  of  the  vena  cava  opening,  we  have  the  hori- 
zontal muscular  fibres  connected  with  the  inferior  ribs  (JS ), 
for  pulling  upon  this  edge  of  the  lumen  simultaneously  during 
inspiration,  and  throwing  the  flood-gate  wide  open  for  afford- 
ing ready  egress  to  the  large  amount  of  blood  flowing  into  the 
lungs  during  this  time,  the  muscular  layer  (o)  upon  the  left 
of  the  middle  leaflet  pulls  also  upon  the  caval  opening.  In  the 
band  of  fibrous  tissue  which  skirts  the  opening  anteriorly  {F, 
N),  we  have  a  needed  brace  for  obviating  strain  from  the  too 
energetic  action  of  the  powerful  muscular  bundles,  especially 
from  the  one  to  the  right  of  the  oesophagus,  which  would  tend 
to  pull  upon  this  portion  unduly,  and  twist  it  upon  itself,  and 
so  disarrange  the  parts  during  inspiration  ;  hence  this  strong 
brace  of  fibrous  tissue.  Nothing  could  be  more  beautiful  than 
these  dispositions  of  force,  with  the  systematic  arrangements 
in  the  structures  for  accomplishing  the  important  ends  in 
view.  It  is  comprehensive  and  superb  mechanics.  The 
ligamentous  arcs,  formed  over  the  psose  and  quadrati  lum- 
borum  muscles,  are  constructed  upon  the  same  principle, 
so  as  not  to  interfere  with  their  functions  or  the  splanchnic 
nerves.    The  ligamentum  arcuatum  internum,  which  is  thrown 


PHYSIOLOGICAL   ANATOMY   IX   VEINS.  243 

across  the  upper  part  of  the  psoas  maguus  muscle  on  either  side, 
is  connected  by  one  end  to  the  outer  side  of  the  body  of  the 
first  (occasionally  of  the  second)  lumbar  vertebra,  and  by  the 
other  to  the  front  of  the  transverse  process  of  the  second  lum- 
bar vertebra ;  while  the  ligamentum  arcuatum  externum, 
thrown  across  the  quadratus  lumborum,  is  attached  by  one 
extremity  to  the  apex  and  lower  margin  of  the  last  rib, 
and  by  the  other  to  the  transverse  process  of  the  second 
lumbar  vercebra,  with  which  the  ligament  for  forming  the 
opening  for  the  splanchnic  nerves  also  connects.  It  is  all  very 
prettily  arranged. 

Adjustments  in  the  Lower  Cava. — We  now  have  to  note 
the  adjustments  in  the  walls  of  the  lower  cava  with  the  action 
in  the  diaphragm.  We  have  seen  that  during  deep  inspira- 
tion the  diaphragm  descends  as  much  as  two  inches ;  hence,  did 
not  some  adjustment  obtain  in  the  walls  of  the  cava,  the  canal 
would  be  bent  upon  itself  during  this  time  and  obstruct  the 
blood,  when  the  purpose  is  to  expedite  the  flow  of  venous  blood 
to  the  lungs,  thereby  bringing  all  the  arrangements  in  the 
diaphragm  to  naught.  On  the  other  hand,  during  forced 
expiration,  and  the  elevation  of  the  diaphragm  in  the  chest 
cavity  which  this  produces,  it  must  inevitably  subject  the  lower 
cava  to  a  degree  of  traction  terrible  even  to  contemplate. 
Besides  it  should  greatly  reduce  the  lumen,  thereby  obstruct- 
ing the  flow  of  blood  were  the  vessel  capable  of  enduring  such 
rude  mechanics,  which  cannot  be  thought  of  for  a  moment  even. 
Now,  then,  for  obviating  these  circumstances,  we  have  the  great- 
increase  which  occurs  in  the  longitudinal  muscles  in  the  walls 
of  the  lower  cava,  adjacent  to  the  diaphragm,  together  with  the 
nerves  for  connecting  it  with  the  solar  plexus,  pneumogastric 
and  phrenic  nerves  for  coordinating  it  with  the  action  in  the 
diaphragm  and  respiration,  so  that  it  contracts  the  longi- 
tudinal axis  during  inspiration,  at  the  same  time  the  lumen  is 
thrown  more  widely  open,  both  within  and  below  the  dia- 
phragm, by  expansion  of  the  transverse  muscles,  the  mus- 
cles in  the  diaphragm  contracting  ;  and  vice  versa  during  ex- 
piration, the  vessel  elongating  pari  passu  with  expansion 
in  the  diaphragm,  thereby  obviating  traction ;  hence,  these 
muscles  and  nerves  to  the  lower  cava  for  coordinating  it  with 


244  PHYSIOLOGICAL   ANATOMY    IN    VI  INS. 

respiration  and   the  action  in  the  diaphragm.      But  in  the 
portion  of    the  cava  above  the   diaphragm,  adjacent  to   the 
heart,  the  muscles  are  absent  for  two  reasons  :   For  the  reason 
that  the  lungs  and  heart  move  downward  during  inspira- 
tion,   and  upward  during  expiration,  moving  together  and 
simultaneously  ;    which  compels  the  upper  cava  and  primitive 
branches  to  elongate  and  contract  in  similar  manner  with  the 
oesophagus  and  lower  cava,  only  the  action  alternates  in  the  two 
localities,  those  above  the  diaphragm  elongating  as  those  below 
it  are  contracting,  and  nice  versa  ;   and  the  anatomical  disposi- 
tions for  effecting  the  actions  are  perfect,  for  the  upper  cava  and 
primitive  branches  are  similarly  endowed  with  the  oesophagus 
and  lower  cava  system,  possessing  both  longitudinal  and  circular 
muscles.    2.  For  the  reason  that  strain  tends  to  fall  in  this  local- 
ity, produced  by  reflux  in  the  right  ventricle  during  the  systole, 
and  the  weight  of  the  fluid  from  above  meeting  the  force  in 
the  diaphragm  from  below,  making  this  the  point  of  resist- 
ance, subjects  the  vessel  to  unusual  pressure,  especially  dur- 
ing expiration,  when  the  blood  tends  to  dam  at  the  right  side 
of  the  heart;  hence  the  dense  fibrous  tissue  in  this  portion 
of  the  lower  cava  for  resisting  strain.     But  we  may  remark, 
en  passant,  that  the  absence  of  the  muscles  does  not  neces- 
sarily imply  a  totally  passive  condition,  since  the  same  deduc- 
tion would  have  to  be  made  for  the  entire  venous  and  arterial 
systems,  by  reason  of  the  intima,  or  elastic  inner  coat,  and  the 
fibrous  and  elastic  ti  sues  in  the  outer  coat,  only  that  muscles 
are  essential  for  increasing  force  and  producing  energetic 
action.     Finally,  this  disposition  of  the  muscles  in  the  walls 
of  the  lower  cava,  as  the  necessary  physiological  adjustments 
with  the  action  in  the  diaphragm,  would  account  for  a  similar 
arrangement  of  the  muscles  in  the  renal  veins,  since  the  kid- 
neys, by  reason  of  their  lying  against  the  diaphragm,  are  also 
made  to  oscillate  with  respiration,  moving  to  and  fro  upon  the 
soft  cushions  of  fat  in  the  long  axis  of  the  body ;    and  did 
not  a  similar  arrangement  obtain  in  the  renal  veins,  these 
also  must  inevitably  be  seriously  affected  by  the  to-and-fro 
movements ;  hence  the  numbers  of  the  longitudinal  muscles 
and  nerves  in  the  walls  of  the  renal  veins,  together  with  the 
nerves  for  connecting  them  with  the  solar  plexus  and  pneu- 


ELONGATION   AND   CONTRACTION  IN  VESSELS.  245 

mogastric  nerves,  for  coordinating  them  with  respiration,  with 
which  the  renal  functions  are  also  necessarily  connected  ;  and 
looking  thence  to  the  vena  porta  and  its  tributaries  (since  a.11 
the  viscera  with  which  they  connect  oscillate  in  similar  manner 
with  respiration),  we  see  that  this  anatomical  arrangement  is 
maintained  throughout,  the  whole  being  rich  in  longitudinal 
muscles.  But  perhaps  one  of  the  most  striking  illustrations 
of  this  function  in  the  vessels  occurs  in  connection  with  the 
splenic  vein,  which  has  adjustment  with  the  functions  in  the 
stomach.  For  example,  the  spleen  is  attached  to  the  great 
€ul  de  sac  upon  the  left,  with  the  vena  portse  at  the  liver 
(Fig.  87);  consequently  when  the  stomach  is  distended  with 
ingesta,  the  splenic  vein  is  fully  twice  the  length  it  has  in  the 
interim  of  digestion — is  not  pulled  into  extension  by  the  dilat- 
ing stomach,  but  elongates  pari  passu  with  it ;  otherwise, 
there  would  be  prodigious  strain,  ending  in  inevitable  disaster. 

Accordingly,  we  find  the  splenic  vein  exceedingly  rich  in 
longitudinal  muscles  ;  whereas,  the  splenic  artery  adjacent  is 
nearly  destitute,  the  muscles  being  nearly  all  circular.  In 
consequence,  during  the  interim  of  digestion,  when  the  splenic 
vein  is  little  more  than  half  its  length  when  the  stomach  is 
full,  the  artery  is  thrown  into  serpentine  folds,  or  the  form  it 
presents  in  the  dead  body  ;  the  vein  measuring  upon  an  aver- 
age less  than  five  inches,  while  the  artery  approximates  nine 
inches.  But  when  the  stomach  is  full  they  are  both  of  the 
same  length ;  hence,  there  can  be  no  doubt  that  the  great 
elongation  and  contraction  in  the  vein  is  due  to  the  number 
of  the  longitudinal  muscles,  which  corresponds  with  the  ac- 
tion in  the  leech,  with  which  it  is  homologous.  In  the  case 
of  the  renal  artery,  however,  the  matter  is  again  different ; 
here  we  have  longitudinal  fibres  in  abundance,  in  correspond- 
ence with  what  occurs  in  the  renal  vein,  to  enable  the  lateral 
movements  in  respiration,  while  its  powers  of  elongating  and 
contracting  are  readily  inferred  from  what  occurs  in  the  arteria 
dor  sails  penis  during  states  of  erection  and  quiescence  ;  since 
the  arrangement  of  the  muscles  is  fundamentally  the  same  in 
both,  the  vessel  elongating  and  contracting  with  the  penis  by 
means  of  the  longitudinal  muscles. 

Thus,  it  is  perceived  that  adaptation  of  means  to  ends  pro- 


240  ELONGATION   AND   CONTRACTION   IN    VESSELS. 

duces  correspondence  in  the  structures,  whether  it  relate  to 
respiration  or  the  action  in  the  penis ;  but  remote  as  these 
functions  would  appear  to  be  from  each  other,  they  both  sus- 
tain the  same  relation  to  the  organic  laws,  compelling  similar 
arrangements  in  the  structures,  in  order  to  produce  the 
actions,  as  must  appear  obvious.  And  it  is  the  same  in  every 
stage  in  development  and  every  organ  in  the  body,  contrasting 
structure  with  structure,  and  function  with  function  ;  as  also 
in  widely  separated  organs  and  functions  where  similar  ac- 
tions are  called  for,  the  same  laws  applying  universally,  and 
adaptation  of  means  to  ends  must  be  in  correspondence,  in  the 
very  nature  of  things. 


CHAPTER  XI. 

AUTOMATISM   IN  THE   LYMPHATICS   AND    THE    MECHANICS    FOB 
CIRCULATING  LYMPH. 

The  Lymphatic  System  Intercalated  between  the  Arterial  and  Venous  Systems, 
Arising  in  the  Tissues  and  Debouching  in  the  Veins  at  the  Root  oi  the  Neck — 
The  Dual  Functions  in  these  Organs  Concerned  in  Drainage  and  Hsematosis — 
The  Vessels  Connecting  with  Respiratory  Movement  in  the  Tissues,  Expanding 
and  Contracting  with  the  Blood-Capillaries  by  Means  of  the  Nervous  Connections 
Subsisting  between  Them — The  Action  in  the  Larger  Vessels  and  Gland  -Structures — 
Muscles  and  Nerves  to  the  Organs  for  Producmg  the  Actions — The  Action  in  the 
Lacteal  System  and  the  Manner  in  which  it  is  Affected  by  the  Action  in  the  Gut 
for  Compelling  Rapid  Movement  of  the  Fluids  in  the  Vessels — The  Relations  which 
this  Sustains  to  Inspiration  and  the  Action  in  the  Venous  System — Physiological 
Experiment  Proving  that  Inspiration  Pumps  the  Lymph  into  the  Venous  System 
simultaneously  with  the  Portal  and  Hepatic  Blood — Proof  of  Automatism  in  the 
Vessels. 

The  lymphatic  system  is  intercalated  between  the  arterial 
and  venous  systems,  taking  its  rise  in  the  tissue  spaces  and 
interstices  and  emptying  into  the  venous  system  at  the  root  of 
the  neck,  at  the  junction  of  the  subclavian  and  jugular  veins 
(Fig.  J  00).  The  left  trunk,  much  larger  than  the  right,  and  known 
as  "the  thoracic  duct,"  receives  the  chyle  and  the  lymph 
from  the  lower  portions  of  the  trunk  and  inferior  extremities, 
inclusive  of  the  left  upper  half  of  the  body  ;  the  right,  known 
as  "the  right  lymphatic  duct,"  a  small  trunk  of  about  an 
inch  in  length,  receives  the  lymph  from  the  right  side  of  the 
head,  right  arm,  right  side  of  the  body,  inclusive  of  the  con- 
vexity of  the  liver,  right  lung  and  right  side  of  the  heart,  the 
orifice  guarded  by  two  semilunar  valves  fur  preventing  ingress 
of  the  venous  blood  ;  the  left  is  similarly  guarded. 

According  to  Recklinhausen,  the  lymphatics  function  as  a 
drainage  system  for  the  tissues,  the  particles  too  large  to  pass 
the  stomata  in  the  capillaries  making  their  way  through  the 
open  ends  of  the  lymph-channels  with  which  the  tissue-spaces 
communicate,   the  tissue-spaces  being   as  so  many  lymph- 


248  RHYTHMIC   MOVEMENT   IN   LYMPHATICS. 

lacunae.  It  seems 'reasonable  only  that  the  drainage  is  unduly 
interrupted  by  the  numerous  glands  thrown  across  the  chan- 
nels in  which  circulation  is  extremely  difficult,  the  dense 
structure  inhibiting  rapid  movement.  Then,  too,  there  are 
the  thymus  and  thyroid  glands,  supra-renal  capsules,  and 
spleen  of  similar  nature  to  be  accounted  for,  which  are 
inexplicable  by  this  theory.  Let  it  be,  however,  for  Ma- 
ture is  utilitarian  to  the  last  degree,  and  may  combine  two 
schemes — blood-making  with  tissue  drainage — in  this  man- 
ner working  the  old  products  for  a  new  role  in  the  organ- 
ism, in  the  genesis  of  white  corpuscles  for  tissue-structure 
and  other  purposes ;  the  matrix  in  the  gland-structures,  the 
slow  circulation  and  rich  materials  conducing  to  the  nutritive 
processes,  thus  serving  as  the  womb  of  the  tissues,  so  to  speak, 
life  springing  out  of  the  ashes,  and  ready  to  commence  again 
the  labor  of  living.     Mirabile  ! 

Without  entering  into  the  mooted  question  of  the  mode  of 
origin  of  the  lymphatics,  whether  by  a  closed  network  of 
capillaries  (larger  than  blood-capillaries),  or  in  the  hue  wall-less 
spaces  in  the  tissue  interstices  traversed  by  the  fluids  to  and 
from  the  cell-brood  and  blood-capillaries,  matters  not  in  the 
argument.  What  immediately  concerns  us  is  the  force  of  this 
circulation  for  compelling  the  lymph  through  the  channels  in 
the  measure  of  the  physiological  requirements  in  the  tissues 
and  lymphatic  glands,  which  it  has  to  traverse  on  its  way  to 
the  venous  system,  together  with  the  manner  of  coordinating 
the  vessels  with  the  blood-vascular  system,  for  producing  con- 
tinuity in  force  and  a  continuous  current  with  the  blood- 
stream, which  the  scheme  calls  for  •;  at  the  same  time,  main- 
taining automatism  in  the  vessels  and  glands,  which  must  not 
be  lost  sight  of  for  a  single  moment,  all  the  others  depending 
upon  it.  The  mechanics  is  easily  understood.  Thus,  when 
the  blood-capillaries  expand  for  reducing  pressure  within 
themselves  and  increasing  it  in  the  tissue  interstices  (pp.  i7o- 
178),  in  order  to  compel  the  fluids  into  themselves,  the  lymph- 
vessels  also  expand  simultaneously,  the  same  nerves  and  nerve- 
ganglia  acting  upon  all  alike,  so  that  they  get  their  share  of 
the  tissue  fluids.  But,  being  larger  than  blood-capillaries,  the 
action  is  necessarily  slower  and  more  difficult,  but  is  sufficient 


KHYTHMIC   MOVEMENT  IN   LYMPHATICS.  249 

to  determine  the  coarser  particles,  which  were  too  large  to  pass 
the  stomata  in  the  more  energetic  blood-capillaries,  into  them- 
selves, floated  into  them  upon  waters,  the  medinm  of  transpor- 
tation ;  passing  in  through  the  open  lumen  in  the  tissue-spaces 
or  th^  larger  stomata  in  the  capillary  walls,  in  the  absence  of 
these  open  ends  to  the  lymph-vessels  ;  while  during  the  sub- 
sequent contraction  the  valves  obviate  reflux,  so  that  suction- 
force  is  made  effective  upon  the  fluids  in  the  tissue  interstices, 
and  no  reason  presents  why  it  should  not. 

.\nd  the  vessels  being  composed  of  the  same  material  as 
the  blood- capillarieSj  namely,  protoplasmic  substance,  and 
connected  with  them  by  nerves,  the  respiratory  movement  in 
the  tissues  should  affect  both  alike  ;  hence,  it  should  have 
the  effect  of  pumping  the  lymph  into  the  lymph-channels 
simultaneously  with  the  fluids  in  the  blood-capillaries,  the 
same  law  applying  to  both.  In  this  manner,  then,  the 
lymph  passes  into  the  lymph-channels  during  expansion  in 
the  vessels,  and  the  high  pressure  this  produces  in  the  tissue 
interstices,  inclusive,  of  course,  of  the  action  in  the  blood- 
capillaries,  the  fluids  flowing  from  high  to  low  pressure ; 
while  during  the  subsequent  contraction  the  valves  would 
obviate  reflux,  at  the  same  time  pushing  the  fluid  up  the 
channel  toward  the  glands  intercalated  in  the  vessels  by  a 
series  of  rhythmical  expansions  and  contractions  similar  to 
what  occurs  in  the  gullet,  with  which  the  vessels  are  homol- 
ogous, due  allowance  being  made  for  the  special  adaptations 
with  the  functions  in  the  organs.  Moreover,  Heller  *  has 
seen  the  lymph-vessels  in  the  mesentery  of  the  guinea-pig  ex- ' 
pand  and  contract  regularly  and  rhythmically.  Then,  again, 
the  veins  contract  for  pushing  the  blood  toward  the  heart 
and  lungs,  and  no  reason  presents  why  the  lymphatics  may 
not  do  likewise ;  seeing,  also,  that  they  connect  with  the 
vaso-motor  and  respiratory  centres,  the  same  as  the  blood 
vascular  system.  Finally,  the  great  difficulty  in  circulating 
lymph,  by  reason  of  the  glands  intercepting  the  stream,  would 
account  for  the  strength  of  the  vessels,  or  the  number  of  mus- 
cles, elastic  and  fibrous  tissue  in  the  walls  of  the  vessels.  Just 
here  it  were  well  to  give  some  idea  of  the  anatomy  in  the 

*  CM.  Med.  Wiss..  1869.  p.  545. 


250 


PHYSIOLOGICAL     ANATOMY. 


glands  to  show  the  difficulties  which  beset  the  way,  and  the 
necessity  for  this  increase  of  force  for  compelling  circulation 
in  the  measure  of  the  requirements.  In  the  first  place,  the 
glands  are  composed  of  a  congeries  of  lymph-nodes  communi- 
cating with  afferent  and  efferent  vessels  (Fig.  93,/,/,  h).     'J  he 


i— \ 


Fig.  93.— Section  of  a  Small  Lymphatic  Gland,  half  diagrammatically  given,  with  the 
course  of  the  lymph,  a,  The  envelope  ;  b,  septa  between  the  follicles  or  alveoli  of 
the  cortical  portion  ;  c,  system  of  septa  of  the  medullary  portion,  down  to  the  hilus 
of  the  organ  ;  e,  lymph-tubes  of  the  medullary  mass  ;  /,  different  lymphatic  streams 
which  surround  the  follicles,  and  flow  through  the  interstices  of  the  medullary  por- 
tion ;  g,  confluence  of  these  passing  through  the  efferent  vessel  ;  h,  at  the  hilus  of 
the  organ  — Frey . 


Fig.  94. — Follicle  from  the  Lymphatic  Gland  of  a  Dog  in  Vertical  Section,  a,  Reticular 
sustentacular  substance  of  the  more  external  portion  ;  b,  of  the  more  internal,  and  c, 
of  the  most  external  and  finely  webbed  part  on  the  surface  of  the  follicle  ;  d,  origin 
of  a  thick  lymph-tube  ;  e,  the  same  of  a  thinner  one  ;  /,  capsule  ;  g,  septa  ;  k,  division 
of  one  of  the  latter  ;  i,  investing  space  of  the  follicle  with  its  retinacula  ;  h,  vas 
afferens  ;  I,  I,  attachment  of  the  lymph-tubes  to  the  septa — Frey. 


PHYSIOLOGICAL    ANATOMY.  251 

internal  arrangements,  as  far  as  made  out,  are  substantially  as 
follows  :  From  the  investing  capsule,  composed  of  fibrous  con- 
nective-tissue intermixed  with  numerous  smooth  muscular 
fibres,  septa  (5,  c)  corresponding  with  the  number  of  the  lymph- 
nodes  are  given  off,  composed  of  the  same  materials  as  the 
capsule,  and  pass  through  the  gland  (c)  to  the  hilus,  investing 
each  of  the  nodes  and  forming  a  similar  capsule  around  them 
also  (Fig.  94,/,  g),  while  in  the  nodes  themselves  the  numerous 
minute  individual  follicles  of  which  they  are  composed  are 
surrounded  by  reticular  tissue  (Fig.  94,  a,  5,  c),  which  serves  the 
purpose  of  a  capsule  to  them  also  (/),  the  whole  thus  connect- 
ing through  and  through,  since  the  reticular  tissue  of  the  fol- 
licles connects  with  the  septa  {i,  i)  forming  the  sustentacular 
tissue,  the  lymph-tubes  also  connecting  with  the  septa  (Z,  I). 

•'But  no  follicle  is  completely  ensheathed  at  its  under  surface 
in  this  system  of  septa.  On  the  contrary,  either  one  or  several 
gaps,  or  even  wide  deficiencies,  are  left,  through  which  the 
follicular  tissue  comes  into  immediate  contact  with  the  medul- 
lary substance.  In  the  same  way,  the  partitions  passing  in- 
ward between  adjacent  follicles  may  be  interrupted  by  massive 
bridges,  as  it  were,  of  lymphoid  tissue,  by  which  these  are  con- 
nected one  with  another,  the  muscular  fibres  being  present  in 
large  numbers"  (Frey).*  Then  we  have  the  lymph-tube,  with 
its  complicated  system  of  vessels  (Fig.  95,  a),f  traversing  the 
reticular  spaces  beticeen  the  follicles  and  the  investing  reticular 
septa  (d,  d),  so  that  a  continuous  passage  is  effected  by  the 
lymph  at  the  same  time  that  it  is  brought  into  intimate  relations 
with  the  follicles,  for  effecting  interchanges  between  them. 
Finally,  we  have  a  system  of  arterial  and  venous  capillaries 
setting  in  and  out  of  the  gland  in  such  manner  as  to  bring 
the  blood  into  iniimate  relation  with  the  follicles  and  gland- 
tubes  (Figs.  96,  95),  the  gland-tube  itself  possessing  its  own 
system  of  blood-capillaries,  so  that  blood  circulates  freely 
through  the  gland,  and  in  the  measure  of  the  physiological 
requirements,  so  that  every  cell  can  get  its  due  supply  of 
arterial  blood,  as  in  every  other  organ.  Now,  then,  what  can 
be  the  purpose  of  all  this  capsular  investment,  with  the  mus- 
cular fibres  richly  interspersed,  other  than  to  effect  rhythmical 

*  Histology  and  Histochemistry  of  Man,  p.  408.  f  Ibid.,  Fig.  404. 


252 


PHYSIOLOGICAL    ANATOMY. 


changes  of  pressure  in  the  follicles  and  lymph-channels  for 
expediting  circulation  in  connection  with  the  special  functions 
in  the  gland,  seeing  that  muscular  structure  is  for  producing 
force,  here  as  elsewhere  ? 
According  to  /Schwartz,  the  muscles  of  the  capsule  at  the 


Fig.  95.— From  the  Medullary  Substaa  jo  of  the  Inguinal  Gland  of  the  Ox  (after  His) . 
a.  Lymph-tube  with  its'  complicated  system  of  vessels  ;  c,  portion  of  another  :  d, 
septa  ;  b,  retinacula  stretched  between  the  tube  and  the  septa. 


^,JU> 


Fig.  96. — Transverse  Section  through  the  Equator  of  three  Peyer's  Patches  of  the  same 
Animal,    a,  The  capillary  network  ;  b,  of  the  larger  circular  vessels.— Frey. 


Ml'SCLES.TO   LYMPH-GLANDS  AND  TESSELS.  253' 

line  of  junction  of  the  medullary  and  cortical  portions  have  a 
principally  radiating  arrangement,  which  is  precisely  what  is 
needed  for  constringing  the  gland  by  producing  contraction  in. 
the  muscles,  and.  vice  versa,  during  expansion  ;  while  the  septar 
passing  inward  between  the  nodes  with  the  broad  bases  at  the 
capsule  (Fig.  88,  5),  and  splitting  up  into  numerous  septa  and 
coming  together  again  for  embracing  the  follicles  for  applying 
a  special  force  to  them,  which  the  functions  call  for,  consti- 
tutes the  fine  adjustment  in  this  mechanics,  in  correspondence 
with  the  universal  rule,  the  difference  being  only  in  adap- 
tive changes  with  the  functions  in  the  gland,  which  follows  as 
a  matter  of  course.  This,  together  with  the  force  in  the  af- 
ferent and  efferent  vessels,  is  sufficient  to  produce  circulatiou 
through  the  glands,  the  force  in  the  chest  and  arterial  system 
also  applying ;  but  is  not  the  force  in  this  circulation,  being 
too  far  removed  for  overcoming  the  difficulties  in  the  glandu- 
lar circulation,  the  force  being  upon  the  ground  where  the 
work  is  done  in  the  gland-structures  and  intercommunicating 
vessels.  Moreover,  the  vessels  themselves  bear  testimony  to 
this  circumstance  in  the  number  of  the  valves  they  contain, 
which  are  strung  like  beads  (Fig.  9?)  along  the  vessels,  and 
the  number  of  muscles  and  nerves  to  them  for  producing 
action  upon  the  lymph  by  means  of  the  rhythmical  expan- 
sions and  contractions  taking  place  in  them,  and  acting  in 
this  respect  as  so  many  hearts  upon  the  stream,  each  one 
representing  a  little  ventricle  or  cardiac  chamber  for  the  pur- 
pose ;  else  this  also  would  be  meaningless.  Moreover,  Heller 
has  observed  such  movements  in  the  lymphatics  of  the  mesen- 
tery, as  has  already  been  observed. 

Turn  we  now  to  the  lacteal  system  and  the  lymph-channels 
of  the  mesentery  for  further  corroborative  evidence  of  the 
principle  which  applies,  here  as  elsewhere  in  the  organism,, 
individualism  pervading  the  whole  of  it,  and  must  do  so  in 
the  very  nature  of  things,  in  order  that  the  vessels  and  glands 
may  perform  their  functions  and  maintain  existence,  while 
force  is  added  in  the  measure  of  the  physiological  require- 
ments. For  example,  contrast  the  action  in  the  villus  with 
the  special  anatomy  in  the  lymph-channels  in  order  to  see' 
adaptations  of  means  to  ends  beautiful  to  look  upon.     ln_  the: 


254 


MUSCLES    -AND   YALYES   TO    LYMPH-VESSELS. 


first  place,  we  have  the  central  lymph-channel  (Fig.  9  ',  I)  in- 
closed in  muscular  walls  (m),  running  the  entire  length  of  the 
channel,  forming  the  sides  over  which  the  pavement  epithe- 
lium is  spread,  the  muscles  extending  from  the  muscularis 
mucosa?  at  the  base  of  the  villus.  ^ow,  then,  when  the 
muscles  contract,  the  effect  is  to  drive  the  lymph  into  the 
lymph-channels  under  the  muscularis  mucosa?  {rete  amplum) 
(Fig.  99,  &),  the  force  in  the  muscles  compelling  it.     On  the 


Fig.  97. — Showing  the  Number  of  the  Lymph- Valves,  with  the  Local  Dilatations  in 
the  Yessels  in  the  Warm-Bl<x>ded  Animals. — Sappey. 

other  hand,  when  the  muscles  expand,  the  low  pressure  this 
produces  in  the  channel  aspirates  the  lymph  ;  then  another 
contraction  followed  by  another  expansion,  and  so  the  little 
mechanism  runs  like  a  heart  for  pumping  the  lymph  into  the 
lymph-channels  below.  But  to  this  we  must  add  the  force  in 
the  muscular  walls  of  the  gut  for  compelling  the  fluids 
into  the  columnar  epithelial  cells  through  the  sieve-like  open- 
ings upon  the  free  surface ;  thence  through  the  memorana 
propria  (e,  e,  e' )  into  the  interior  of  the  villus,  the  high  pres- 
sure it  produces  in  the  intestine,  together  with  the  actions  in  the 
villus  and  the  epithelia  (for  they  are  all  connected  by  nervous 


MUSCLES   TO   LACTEALS. 


255 


force)  compelling  this  circumstance.  No  wonder,  then,  that 
fat  or  anything  else  absorbable  should  be  rapidly  absorbed. 
And  there  is  no  necessity  for  endowing  any  portion  of  the 
mechanism  with  extraordinary  powers  in  order  to  account  for 
the  circumstance,  such  as  that  the  epithelial  cells  act  like 


#■  Tiff  i 


Fig.  98. — Diagrammatic  Section  of  a  Villus. — Watney.  ep,  Epithelium  only  partially 
shaded  in  ;  I,  central  chyle-vessel — the  cells  forming  the  vessel  have  been  less  shaded 
to  distinguish  them  from  the  cells  of  the  parenchyma  of  the  villus  ;  m,  muscle  fibres 
running  up  by  the  side  of  the  chyle- vessel.  It  will  be  noticed  that  each  muscle  fibre 
is  surrounded  by  the  reticulum,  and  by  this  reticulum  the  muscles  are  attached  to 
the  cells  forming  the  membrana  propria,  as  at  e',  or  to  the  reticulum  of  the  villus. 
Ic,  Lymph  corpuscles,  marked  by  a  spherical  nucleus  and  a  clear  zone  of  protoplasm  ; 
I',  upper  limit  of  the  chyle-vessel ;  e,  e,  e',  cells  forming  the  membrana  propria.  It 
will  be  seen  that  there  is  hardly  any  difference  between  the  cells  of  the  parenchyma, 
the  endothelium  of  the  upper  part  of  the  chyle-vessel,  and  the  cells  of  the  membrana 
propria,  v,  Blood-vessels  ;  z,  dark  hue  at  the  base  of  the  epithelium  formed  by  the 
reticulum.  It  will  be  seen  that  the  reticulum  penetrates  between  all  the  other  ele- 
ments of  the  villus.  The  reticulum  contains  thickenings  or  "nodal  points."  The 
diagram  shows  that  the  cells  of  the  upper  part  of  the  villus  are  larger  and  contain  a 
larger  zone  of  protoplasm  than  those  of  the  lower  part.  The  cells  of  the  upper  part 
of  the  chyle-vessel  differ  somewhat  from  those  of  the  lower  part,  in  that  they  more 
nearly  resemble  the  cells  of  the  parenchyma. 


256  MUSCLES  TO   LACTEALS. 

amoeba)  and  eat  the  fat.*  But  in  my  heart  of  hearts  I  thank 
the  distinguished  author  for  the  large  individualism  in  the 
cells  and  tissues  it  shadows  forth,  and  he,  of  all  men,  is 
closest  to  the  mechanics  in  the  animal  organism.  But  the 
amoeba  must  yield  up  some  of  its  license  in  the  compound 
organism  for  the  common  weal,  and  must  take  a  subordinate 
position  in  blending  with  the  common  functions  with  which 
their  actions  are  coordinated.  And  in  the  case  before  us,  it  is 
not  compelled  to  extend  branched  processes  into  the  canal  for 
embracing  the  food,  but  only  to  expand  the  oral  orifices  or 
sieve-like  openings,  while  the  food  is  pushed  into  them  under 
the  force  in  the  gut,  the  body  of  the  cell  expanding  in  order 
to  receive  it ;  in  consequence,  the  epithelia  rapidly  become 
double  their  size,  and  of  a  milky  appearance  from  the  amount 
of  fat  and  other  aliment  they  contain.  And  if  Professor  Foster 
meant  this,  we  are  in  full  accord;  but  I  do  not  think  he  included 
the  pressure  in  the  gut  for  assisting  the  action. 

We  have  but  to  follow  the  lymph  to  the  channels  be- 
neath the  muscu1  arts  mucosa  to  see  how  the  force  in  the  gut 
applies  to  this  also,  together  with  the  force  in  the  numerous 
muscles  forming  the  walls  of  the  lymph-vessels,  by  means  of 
which  the  local  action  is  increased  and  the  stream  regulated 
with  the  capacity  in  the  mesenteric  glands,  with  which,  of 
course,  it  must  have  adjustment ;  otherwise,  choking  and 
engorgement  would  follow  as  an  inevitable  sequence  of  excess- 
ive action  in  the  lymph-vessels.  The  following  beautiful 
illustration, f  from  Teichmann,  on  the  lymphatic  system,  will 
serve  for  impressing  the  matter.  For  example,  the  lymph- 
channel  (there  are  several  given  here)  in  the  villus  debouch 
into  the  close-meshed  lymph-vessels  {rete  cuu/ustum)  be- 
neath the  muscularis  mucosce  (not  given  in  the  picture)  at  the 
base  of  the  villi  (a,  b),  which  rapidly  expand  into  great  lymph- 
chambers  (c),  in  contact  with  the  muscular  cylinder  (g,  h), 
through  which  the  efferent  vessels  (d,  d)  pass  to  get  beneath 
the  peritoneal  layer  of  the  gut  in  order  to  reach  the  mesen- 
teric glands,  situated  in  the  folds  of  the  mesenteric  ligament. 

*  A  Text-Book  of  Physiology,  p.  819.     M.  Foster,  M.  A.,  M.  D.,  F.  R.  S.,  Prae- 
lector  in  Physiology  and  Fellow  of  Trinity  College.  Cambridge, 
f  Ludwig  Teichmann,  "  Das  Saugader  System."    Leipzig.    1861. 


MUSCULAR  FORCE  FOR  THE  LYMPH.  257 

Finally,  that  the  lymph-vessels  and  chambers  are  muscu- 
lar organs  provided  with  valves  for  obviating  reflux,  and 
furnished  with  nerves  from  the  adjacent  ganglionic  layers 
(Meissner's  and  Auerbach's,  which  we  will  come  to  presently) 
for  producing  the  rhythmical  expansions  and  contractions 
which  occur  in  them,  and  which  also  connect  with  the  great 
pumping  movement  in  the  muscular  cylinder  itself,  the  nerves 


Fig.  99. — Perpendicular  Section  through  one  of  Peyer's  Patches  in  the  lower  part  of  the 
Ileum  of  the  Sheep. — Teichniann.  a,  a,  Lacteal  vessels  in  the  villi ;  6,  6,  superficial 
layer  of  the  lacteal  vessels  (rete  angustum)  ;  c,  c,  deep  layer  of  the  lacteals  (rete 
amplum)  ;  d,  d,  efferent  vessels  provided  with  valves  ;  e,  LieberMihn's  erlands  ;  /, 
Peyer's  glands  ;  g,  circular  muscular  layer  of  the  wall  of  intestine ;  h,  longitudinal 
laver  ;  i,  peritoneal  layer. 

connecting  through  and  through,  the  same  as  in  the  case 
of  the  villus,  gland-structures  and  blood-vessels  for  produc- 
ing simultaneous  action  throughout,  the  principle  being  the 
same.  Bur  once  within  this  system  of  reservoirs,  it  is  easy  to 
perceive  that  the  force  in  the  gut  is  available  for  compelling 
the  lymph  through  the  channels,  and  no  power  can  prevent 
it  but  paralysis  in  the  gut  itself,  as  must  appear  obvious.  In 
other  words,  every  contraction  of  the  muscular  cylinder  by 


258  MUSCULAR   FOKCE   FOR   THE   LYMPH. 

compressing  the  air  and  aliment  firmly  against  the  mucous 
membrane  and  submucosa,  in  which  the  lymph-vessels  and 
reservoirs  are  situated,  thence  against  the  resistent  muscular 
cylinder,  whence  there  is  no  escape,  should  have  the  effect  of 
constringing  the  organs,  closing  upon  them  like  the  hand  for 
the  purpose .  Please  look  at  the  anatomy  well.  No  mechanics  in 
the  body  exceeds  it  for  simplicity  and  effectiveness.  It  is  a  work 
of  the  gods.  By  the  action  in  the  gut,  then,  and  the  action  in  the 
lymph-vessels  and  reservoirs,  which  are  coordinated  with  the 
muscular  cylinder,  it  is  passed  by  the  efferent  vessels  (d,  d) 
through  the  gut,  thence  through  the  vessels  to  the  glands  in 
the  mesentery,  by  means  of  rhythmical  expansions  and  contrac- 
tions in  the  vessels,  the  same  as  in  the  systemic  lymph-vessels, 
and  which  would'  include  the  action  in  the  glands,  and  there- 
fore need  not  detain  us.  Finally,  to  this  we  must  add  the  force 
in  the  walls  of  the  abdomen  and  the  pumping  action  in  respi- 
ration, the  same  as  in  the  venous  system,  into  which  it  empties, 
for  producing  simultaneous  action  and  correspondence  in  the 
currents,  which  the  scheme  calls  for,  the  same  mechanics  apply- 
ing to  both  alike.  In  order  to  definitely  ascertain  this  circum- 
stance, also,  I  performed  the  same  experiment  as  in  the  case  of 
the  portal  blood  and  liver  circulation,  taking  the  body  apart 
in  the  same  way,  carefully  ligating  the  vessels  before  dividing 
them  ;  also  the  vena  cava  and  oesophagus.  And  everything 
being  ready,  the  thoracic  duct  was  snipped,  when  the  lymph 
spurted  out  with  force  to  some  feet  distant ;  and  not  waiting 
for  the  vessels  to  empty  themselves  ;  I  bore  down  upon  the 
diaphragm  with  my  open  hand  in  imitation  of  inspiration, 
and  every  time  I  did  this,  the  stream  rose  instantaneously 
into  a  jet  the  same  as  the  venous  blood  had  done.  So,  then, 
there  can  be  no  earthly  doubt  that  inspiration  pumps  the  lymph 
and  venous  blood  simultaneously  into  the  chest-cavity  and 
lungs,  the  amount  being  determined  by  the  energy  and  depth  of 
inspiration  and  the  fullness  of  the  vessels.  The  following 
beautiful  diagram  of  the  lymphatic  system  (Fig.  100),  by  Pro- 
fessor Dal  ton,  will  serve  for  impressing  the  matter.  The 
lacteal  system,  spread  out  in  the  vast  mucous  surface,  acted 
upon  by  the  muscular  cylinder  and  the  muscles  in  the  lymph- 
channels,  this  supplemented  by  the  force  in  the  walls  of  the 


THE   FORCE    IIST   THE   ABDOMEN.  259 

abdomenfor  compressingthegut,  mesenteric  vessels,  and  glands 
during  inspiration,  lifts  the  lymph  to  the  lungs  the  same  as  the 
portal  and  hepatic  blood  and  the  blood  in  the  lower  cava  sys- 
tem, and  simultaneous  with  them.     Please  look  at  it.     I  do 


Pig.  100. — Diagrammatic  Representation  of  the  Lymphatic  System. — Dalton. 

not  see  necessity  for  adding  another  word,  the  matter  being  so 
very  obvious. 

In  conclusion :  That  there  is  automatism  in  the  lymph  ves- 
sels is  not  only  logically  true,  but  the  fact  is  scientifically 
ascertained.  Notably,  in  batrachia  four  lymph-hearts  pre- 
sent, one  behind  each  femoral  joint,  for  pumping  the  lymph 


260  PROOF    OF   AUTOMATISM. 

coming  from  the  hinder  portions  of  the  body  into  the  ischiadic 
veins,  and  one  in  front  of  each  scapula  for  pumping  the  lymph 
from  the  anterior  portions  into  the  jugular  veins,  the  organs 
expanding  and  contracting  regularly  and  rhythmically  from 
fifty  to  sixty  times  per  minute.  Moreover,  they  are  not  syn- 
chronous with  the  action  in  the  heart  or  with  respiration,  or 
even  with  one  another,  possessing  independent  action. 

' k  In  the  large  ceratoplirys  cornuta  two  pairs  of  ischiadic 
lymph-hearts  have  been  found.  In  the  tortoise  the  pelvic 
lymph-hearts  are  two,  of  a  more  circumscribed,  rounded  form, 
situated  on  each  side  of  the  bodies  of  the  vertebrae,  between  the 
femoral  joints  and  the  hind-border  of  the  carapace ;  the  valves 
at  the  inlets  and  outlets  of  the  lymph-conduits,  impressing  the 
course  of  motion  of  the  fluid,  are  here  readily  seen." 

' '  In  Lizards  and  Crocodiles  the  pelvic  lymph-hearts  are  situ- 
ated near  or  upon  the  diapophyses  of  the  first  caudal  vertebra. 
In  Pseudopus  P alias  ii  they  lie  between  the  muscles  ivpon  the 
sacral  diapophyses,  receiving  the  lymph  each  by  a  single  con- 
duit from  the  great  abdominal  sinus,  and  transmitting  it  to 
the  umbilical  veins  ;  they  pulsate  about  fifty  times  in  tlie 
minute.  In  true  serpents  (Python,  e.  r/.)the  lymph-hearts  are 
elongate,  and  situated  behind  the  last  pair  of  ribs  and  upon 
the  rib-like  diapophyses  of  the  anterior  caudal  vertebrae  ;  they 
receive  the  lymph  by  three  orifices  at  one  end,  and  transmit  it 
by  two  opposite  orifices  to  conduits  communicating  with  the 
caudal  vein.  The  three  tunics  of  these  hearts,  of  which  the 
middle  one  is  muscular,  with  the  inferent  and  afferent  valvular 
-v' ' ruetures,  are  well  displayed  in  python"  (Owen).*  Italics  are 
added. 

The  following  from  Hermann  possesses  renewed  interest : 

"In  amphibia  and  certain  birds  (ratitse),  the  movement  of  the 
lymph  is  assisted  by  the  rhythmical  pulsation  of  lymph-hearts, 
of  which  four  exist  in  the  frog,  two  in  other  amphibia,  and 
one  in  the  ostrich.  The  central  nervous  organ  connected  with 
this  rhythmical  motion  is  said  by  some  to  be  in  the  spinal 
cord,  and  by  others  in  the  hearts  themselves.  In  guinea-pigs 
a  rhythmical  contraction  has  recently  been  observed  (A.  Hel- 

*  Comparative  Anatomy  and  Physiology  of  Vertebrates,  vol.  I.,  pp.  459-460. 


PEOOF    OF   AUTOMATISM.  26 J 

ler)  in  the  lymphatics  of  the  mesentery.  As  this  proceeds 
along  those  portions  of  the  vessel  between  the  valves  with  a 
regular  progression  toward  the  larger  trunks,  it  must  be  re- 
garded as  a  species  of  cardiac  mechanism. 

"In  the  frog,  the  rhythmical  movement  of  the  lymph-heart 
seems  to  be  dependent  upon  the  integrity  of  spinal  centres 
(Volkmann),  one  situated  opposite  the  third  vertebra  for  the 
anterior  pair,  and  one  opposite  the  sixth  vertebra  for  the  pos- 
terior pair.  It  is,  however,  affirmed  (Eckhard,  Schiif,  Goltz, 
"Waldeyer)  that  the  power  of  regular  movement  may  some- 
times be  recovered  after  it  has  been  abolished  by  dividing 
the  spinal  nerves,  supplying  the  hearts,  or  by  destroying  the 
spinal  cord. 

"  The  lymph-hearts  are.  further,  constantly  inhibited  from 
the  optic  lobes  (Suslowa),  this  inhibition  being  controlled  by 
centripetal  stimuli,  proceeding  from  skin  or  intestines  (Goltz)  " 

The  following  is  from  Gegenbaur  :* 

"  The  lynrphatic  trunks  of  birds  have  the  same  characters,  but 
in  them  both  the  large  trunk  in  front  of  the  aorta  (thoracic 
duct)  and  the  small  vessels  are  more  independent.  As  in  the 
reptilia,  the  thoracic  duct  opens  into  the  superior  vense  cavae 
(vense  brachiocephalicse).  At  the  commencement  of  the  tail 
the  lymphatic  system  is  also  connected  with  the  ischiadic 
veins,  or  with  the  afferent  renals,  in  which  point  they  resemble 
the  amphibia  and  reptilia.  In  the  mammalia,  the  walls  of 
the  lymphatic  system  are  still  more  differentiated,  although 
it  often  happens  that  in  them  also  the  sheath  of  the  arteries 
bounds  the  course  of  part  of  the  lymphatic  current.  Where 
they  do  not  accompany  the  blood-vessels,  they  form  frequent 
anastamoses,  or  wide-meshed  plexuses,  and  are  distinguished 
by  valves,  as  are  the  same  parts  in  birds.  The  lymphatic 
vessels  of  the  hinder  extremities,  as  well  as  the  chyle-ducts, 
unite  into  a  chief  trunk  in  the  abdomen,  which  is  rarely 
paired,  and  the  origin  of  which  is  frequently  distinguished 
by  a  considerable  enlargement  (cisterna  chyli).  Thence  they 
are  continued  into  a  thoracic  duct,  which  opens  into  the 
commencement  of  the  left  brachio-cephalic  vein ;   the  trunks 

*  Elements  of  Comparative  Anatomy,  p.  595. 


262  PROOF    OF   AUTOMATISM. 

of  the  lymphatics  of  the  anterior  parts  of  the  body  (of  the 
head  and  anterior  extremities),  and  of  the  wall  of  the  thorax, 
open  into  and  on  either  side  of  the  same  vein."  Italics  are 
added.  Thus,  with  progress  in  development  the  lymph-hearts 
disappear,  but  in  lieu  of  them  we  have  increasing  differentiation 
in  this  system  of  vessels,  which  become  more  and  more  muscu- 
lar ;  and  in  place  of  the  local  dilatations  forming  the  so-called 
lymph-hearts,  which  contain  three  coats,  the  middle  one 
muscular,  in  the  warm-blooded  animals  the  muscular  coat 
extends  over  the  entire  walls  of  the  vessels,  the  same  as  in  the 
veins  with  which  they  are  homologous,  only  that  the  valves  are 
greatly  increased  in  number  and  the  external  fibrous  tunic  is 
thicker  and  firmer,  which  more  than  compensates  for  the 
k' hearts"  by  greatly  increasing  the  force  in  the  vessels  which 
it  effects.  At  the  same  time,  it  is  seen  that  automatism  is 
maintained,  and  must  be  so  in  the  very  nature  of  things. 


CHAPTER  XII. 

NERVES    TO    THE    VISCERA    IN    THE    ABDOMEN,    AID    MODE    OF 
CONNECTING  THEM   WITH   THE   CEREBRO-SPIiNAL   AXIS. 

The  Double  Ganglionic  Dorsal  Chain  in  Vertebrates  the  Analogue  of  the  Double 
Ganglionic  Chain  in  the  "Worms — Nerves  of  Meissner — Nerves  of  Auerbach — Mode 
of  Connecting  them  with  the  Solar  Plexus  and  Central  Nervous  System — Relations 
of  the  Nerves  of  Meissner  and  Auerbach  with  the  Intestinal  Mucous  Membrane  and 
Epithelium — Mode  of  Controlling  the  Blood-Supply  from  the  Aorta-Trunk  by  Means 
of  the  Coeliac  Axis,  Superior  and  Inferior  Mesenteric  Arteries  and  then'  Branches,  so 
that  Every  Organ  and  Fractional  Portion  of  the  same  can  Regulate  their  own  Sup- 
plies in  the  Measure  of  the  Functional  Activities — Connection  of  the  Pneuniogastric 
Nerves  with  the  Solar  Plexus — Mode  of  Connecting  the  Solar  Plexus  and  Spinal 
Ganglia  with  the  Dorsal  Nerves  and  Spinal  Medulla,  the  Nerves  of  the  Ganglionic 
Chain  running  up  Both  Roots  of  the  Spinal  Nerves  to  reach  the  Spinal  Medulla — 
Every  Nervous  Ganglion  a  Centre  of  Nervous  Force,  Possessing  both  Sets  of  Fibres, 
or  Dilator  and  Contractor  Nerves — The  Manner  Reflex  Action  is  Produced  in  the 
Spinal  Cord  and  Medulla  Oblongata  through  Sensory  Impressions  in  the  Mucous 
Membrane  and  Cutaneous  Surface,  for  Expanding  and  Contracting  the  Vessels 
and  Maintaining  a  Balance  in  the  Circulation — Relative  Amount  of  Nervous  Force 
sent  to  the  Viscera  through  the  Pneumogastric  and  Splanchnic  Nerves,  Illus- 
trated by  a  Case  of  Fracture  of  the  Fourth  and  Fifth  Cervical  Vertebras,  Producing 
Displacement,  with  Laceration  and  Compression  of  the  Spinal  Cord — Effects  from. 
Division  of  the  Pneumogastric  Nerves  in  the  Neck,  upon  the  Lungs,  Stomach  and 
Intestines — Death  by  Thirst  and  Starvation,  Precipitated  by  Pulmonary  Congestion 
from  Paresis  in  the  Capillary  Network — The  Action  in  the  Kidneys,  and  the  Rela- 
tions they  Sustain  to  the  Solar  Plexus  and  Spinal  Axis  through  Intercom  muni  eating 
Nerves — Ditto  Ureters  and  Urinary  Bladder. 

It  will  now  be  necessary  to  briefly  refer  to  the  nervons  ap- 
paratus for  producing  the  movements  in  the  intestines,  and 
for  coordinating  them  with  respiration,  which,  of  course, 
would  include  the  glandular  appendages  with  the  portal  and 
lacteal  systems  for  producing  correspondence  throughout, 
which  the  scheme  calls  for.  Separate  action  in  the  alimentary 
canal  is  well  enough  for  maintaining  automatism  in  connec- 
tion with  the  special  functions  in  the  organs ;  but  since  the 
whole  relates  to  the  cell-brood  in  the  tissues,  it  is  manifest 
that  this  involves  correlation  of  the  nervous  forces  with  the 
federal  centre  for  the  organism  for  compelling  response  to  the 


264  CORRELATION   OF   NERVOUS   FORCE. 

wants  in  the  cell-brood,  which  is  effected  by  means  of  the 
double  rows  of  spinal  ganglia  and  pneumogastric  nerves  which 
converge  in  the  solar  plexus,  the  common  nervous  centre  for 
the  viscera,  so  that  the  whole  performs  as  but  a  single  organ, 
only,  under  the  action  of  the  nervous  forces  radiating  from  the 
medulla  oblongata  and  respiratory  centre,  in  harmonious  con- 
cert with  the  action  in  the  lungs,  with  which  everything  is  co- 
ordinated, to  the  end  that  a  balance  may  be  maintained  in  the 
organism;  otherwise  impossible.  Now,  then,  with  reference  to 
this  double  ganglionic  chain  extending  the  length  of  the  spinal 
column,  the  analogue  of  the  double  ganglionic  chain  in  the 
worms,  as  it  were,  a  remnant  of  this,  which  the  principle  in  the 
mechanics  compels  to  be  retained,  the  structures  in  the  gut  be- 
ing fundamentally  the  same  as  in  the  worms,  as  before  re- 
marked. And  the  gut  being  folded  upon  itself  in  the  central 
portions,  in  form  of  the  mesentery,  which  the  shortness  of  the 
animal  compels,  the  ganglionic  chain  necessarily  undergoes  a 
similar  folding,  whereby  the  solar  plexus  is  formed  ;  while  con- 
tinuity with  the  basal  cerebral  ganglia  is  maintained  in  the  man- 
ner as  stated,  or  by  means  of  the  trisplanchnic  and  pneumogas- 
tric nerves  ;  the  former  being  thickened  and  elongated  links,  re- 
sulting from  the  folding  up  of  the  chain,  the  latter  an  adap- 
tation to  the  changes  in  the  intestines  which  are  now  detached 
from  the  muscular  envelope,  compelling  this  mode  of  nerve- 
distribution  for  effecting  coordination  with  the  containing 
wails,  as  has  already  been  fully  set  forth.  In  this  manner, 
then,  we  have  the  massing  of  the  ganglia  in  the  solar  plexus 
readily  accounted  for,  and  which  would  include  all  the  other 
arrangements  that  obtain  respecting  it,  as  we  shall  see  further 
on.  The  spinal  axis  is  an  outgrowth  of  the  basal  cerebral  gan- 
glia, and  possessing  separate  reflex  nervous  centres  connecting 
with  the  viscera,  vascular  system,  and  the  voluntary  motor 
apparatus,  represented  by  the  gray  central  portions,  subserves 
important  functions  in  the  mechanics  for  maintaining  the  local 
actions  and  enabling  ready  coordination  with  the  federal  centre 
in  the  medulla  oblongata. 

Commencing  with  the  mucous  surface,  we  have,  then,  the 
following  separate  lines  of  nervous  ganglia,  with  the  interven- 
ing nerves,  for  producing  the  local  actions  in  the  intestines 


CORRELATION   OF   NERVOUS   FORCE.  265 

and  for  connecting  them  with  the  central  nervous  system, 
namely :  1.  The  thick  ganglionic  layer  of  nervous  ganglia  in 
the  sub  mucosa,  or  nerves  of  Meissner  (Fig.  107,  1),  with  the 
nerves  running  thence  into  the  villi  to  connect  with  the  epithe- 
lium (2),  which  includes  the  whole  mucous  surface,  at  the  same 
time  supplying  the  capillaries  and  muscularis  mucosae. 

2.  Next  to  this  ganglionic  layer,  or  between  the  circular  and 
longitudinal  muscles,  is  the  thick  ganglionic  layer  of  Auerbach 
(Fig.  107,  4),  for  supplying  the  muscular  cylinder,  at  the  same 
time  giving  off  inter- communicating  nerves  (3)  to  the  gan- 
glionic layer  of  Meissner,  which  also  gives  off  inter-communi- 
cating nerves,  whereby  the  muscular  cylinder  and  the  mucous 
membrane  are  fully  coordinated,  so  that  it  all  works  together 
harmoniously — vascular  apparatus,  glandular  apparatus  and 
muscular  cylinder,  inclusive  of  the  muscularis  mucosse,  and 
which  includes  the  stomach  as  well,  since  the  nervous  arrange- 
ments are  fundamentally  the  same  in  it. 

3.  Next  to  this  ganglionic  layer,  but  outside  the  gut  and 
removed  to  a  distance  from  it,  we  have  the  federal  centre  for 
the  intestines  in  the  great  solar  plexus  (Fig.  109),  with  the 
nerves  extending  thence  over  the  superior  mesenteric  artery 
(Fig.  108,  1,  2)  to  the  intestines  to  connect  with  the  other  two 
layers  (Fig.  107,  5,  5),  whereby  the  whole  intestinal  apparatus, 
inclusive  of  the  glandular  appendages,  is  coordinated  and 
force  is  increased  in  the  gut ;  the  same  remark  applying  to  the 
stomach,  the  nerves  extending  to  it  and  the  glandular  ap- 
pendages over  the  branches  of  the  cceliac  axis  (Fig.  109,  4). 

4.  Finally,  we  have  the  solar  plexus  connected  with  the ' 
central  nervous  system  by  means  of  the  sj)lanchnic  and  pneu- 
mogastric  nerves  (Fig.  109,  2,  3),  as  before  remarked.  In  this 
manner,  then,  the  whole  is  connected  through  and  through 
with  the  cerebro-spinal  axis  for  coordinating  the  viscera  with 
respiration,  and  for  increasing  nervous  force  as  occasion  may 
require.  It  is  comprehensive,  but  the  continuity  of  relation 
effected  by  means  of  the  nerves  for  producing  the  actions 
with  respiration  is  easily  seen  and  readily  understood. 

In  order  to  perfect  the  mental  picture  of  the  nervous  appa- 
ratus, however,  it  will  be  necessary  to  enter  a  little  more  into 
the  minutiae.     While  the  two  ganglionic  layers  in  the  walls  of 


266  NERVES   TO   THE   COLUMNAR   EPITHELIUM. 

the  intestines  are  plain  enough,  and  minutely  described,  unfor- 
tunately this  cannot  be  said  of  the  nerves  to  the  villi  and 
columnar  epithelium,  which  are  not  yet  definitely  ascertained  ; 
and  we  are  compelled  to  have  recourse  to  analogous  structures  •, 
notably,  the  columnar  cells  of  the  salivary  glands,  in  order 
to  fully  complete  the  nervous  connections  that  obtain  in  the 
intestines.  According  to  the  exhaustive  researches  of  Pniiger, 
the  columnar  cells  sustain  the  most  intimate  relations  to 
the  nerves,  which  blend  with  them  and  form,  so  to  speak,  the 
footstalk  of  the  cells  (Figs.  101,  102),  the  nuclei  of  the  columnar 
cells  being  evolved,  as  it  were,  in  the  terminal  extremity  of 
the  nerves  (Fig.  103).  The  following  extensive  excerpt*  will 
place  the  matter  fully  before  the  reader  : 

' '  When  we  see  the  axis  cylinder  and  its  fibrils  to  be  directly 
continuous  with  the  fibrils  of  the  columnar  cells,  without  any 
difference  being  perceptible  between  the  axis  cylinder  and  the 
fibrils  of  these  cells,  we  may  legitimately  describe  the  nerve  as 
extending  to  the  point  where  it  joins  the  substance  of  the  body 
of  the  cell.  That  is  the  most  natural  explanation  that  can  be 
given.  This  explanation,  however,  possesses  the  greatest  sig- 
nificance in  regard  to  the  mode  of  development  of  the  glandular 
epithelium,  because  it  directly  follows  that  the  young  nuclei 
originate  in  the  axis  cylinders,  and  that  the  gland  cells,  which 
at  a  later  period  seem  to  constitute  a  thickening  of  the  axis 
cylinder,  bud  forth,  as  it  were,  from  the  nerves.  This  expla- 
nation renders  it  intelligible  why  the  nuclei  of  the  columnar 
cells  are  so  indifferent  during  the  multiplication  of  the  epithe- 
lium. In  opposition  to  this  view,  which  I  regard  as  the  most 
probable,  it  may  be  urged  that,  in  consequence  of  the  intimate 
fusion  of  nerve  substance  and  epithelium  at  the  periphery,  no 
sharp  limit  can  be  drawn,  showing  where  the  one  ceases  and 
the  other  begins ;  and  that,  moreover,  it  is  probable  that 
imperceptibly  fine  processes  are  given  off  by  the  nucleus  of  the 
columnar  epithelial  cells,  which  become  detached  at  an  early 
period  by  fission.  That  the  nuclei  of  the  salivary  cells  have 
processes  cannot,  however,  be  regarded  as  forming  a  valid 
objection  to  my  view,  since  the  young  nuclei  may  really  be 
thickenings  of  the  axis  cylinder  fibrils. 

The  Salivary  Glands.     Strieker's  Manual  of  Histology,  p.  314. 


NERVES   TO   THE   COLUMNAR  EPITHELIUM. 


267 


"I  may  further  adduce,  as  a  weighty  argument  in  favor  of 
my  view,  that  the  fibrils  of  the  axis  cylinder  do  not  terminate 
at  the  surface  of  the  fully  developed  salivary  cells,  but,  as 
in  the  case  of  the  ganglion  cells,  may  be  traced  into  their  very 
substance. 

"Now,  since  the  finest  axis  cylinders  and  fibrils  extend  to  the 
columnar  epithelial  cells,  and  are  connected  with  the  processes 


Pig.  101.—  Termination  of  Medullated  Fibres  Treated  with  Perosmic  Acid  in  Isolated 
Salivary  Cells.  A,  thick  branched  fibres  distributed  to  large  salivary  cells  ;  B,  fine 
nerves  distributed  to  smaller  salivary  cells.  From  the  submaxillary  gland  of  the 
Rabbit.    Magnified  590  diameters. — Pfiiiger. 


Fig.  102.— A,  B,  multipolar  cells  in  connection  with  salivary  cells.  Magnified,  A  480, 
B  500  diameters.  C,  peculiar  cells  with  round  thick  processes,  and  containing 
refractile  fat  particles.    Magnified  590  diameters.— Pfiiiger. 

that  are  in  course  of  development,  and  since  portions  of  these 
processes  subsequently  become  large  salivary  cells,  connected 
with  thick  medullated  nerve  fibres,  it  follows  that  the  nerves 
must  increase  coincidently  with  the  young  epithelium  to  which 
they  belong.  Among  these  metamorphoses  there  also  occurs 
a  mode  of  termination  of  the  medullated  nerves,  to  which  I 
some  time  ago  called  attention,  and  which  consists  in  the  nerve 


268  NEKVES   TO   THE   COLUMNAR   EPITHELIUM. 

suddenly  undergoing  frequent  division,  then  enlarging,  and 
containing  finely  granular  protoplasm,  with  many  nuclei  of 
various  sizes.  I  have  named  this  mode  of  nerve  termination, 
that  by  a  'protoplasmic  foot.'  If,  as  I  have  sometimes  ob- 
served, many  of  the  nuclei  appear  to  be  provided  with  fibres, 
which  can  be  followed  into  the  interior  of  the  nerve  fibres,  it  is 
highly  suggestive  of  the  development  of  the  gland  cells  from 
the  nerves. 

"In  regard  to  every  explanation,  it  must  be  observed  that 
transitional  forms  may  occur,  respecting  which  it  is  impossible 
to  say  whether  they  are  epithelial  or  nervous.  The  continu- 
ous and  luxuriant  neoplastic  formation  taking  place  in  the 
substance  of  the  salivary  ducts  presupposes  their  regenera- 
tion, respecting  which  I  have  formed  my  own  opinion,  but 
have  arrived  at  no  definite  conclusion.  In  like  manner  the 
persistent  neoplastic  formation  of  the  alveoli  in  adult  animals 
determines  an  atrophic  detachment  of  those  already  present 
In  moles  I  have  sometimes  found  the  alveoli  with  pale  off- 
shoots of  various  forms,  and  pale  finely  granular  contents, 
which  may  be  such  atrophied  and  separated  alveolar  seg- 
ments. 

"  I  first  comprehended  the  complexity  of  all  forms  of  salivary 
glands  when  I  recognized  the  constant  production  and  disinte- 
gration taking  place  in  them,  which  is  referable  to  the  nerve 
substance." 

This  genesis  of  the  columnar  cells  in  nerve- extremities  may 
be  seen  in  the  different  stages  of  cell  growth  in  the  follow- 
ing cut  (Fig.  103,  a,  b,  c,  d,  e),  beginning  with  the  nuclei  and 
progressing  till  the  cell  is  perfected  {E)  Accordingly,  I  have 
ventured  to  connect  the  columnar  cells  of  the  villi  with  the 
nerves  proceeding  from  the  ganglionic  layer  of  Meissner 
(Fig.  107,  1)  to  the  villi  (2)  This  portion  of  the  picture,  there- 
fore, is  ideal.  But  considering  the  energy  in  the  absorptive 
processes  and  the  rapidity  with  which  the  cells  expand,  and 
especially  the  quick  response  to  stimulus  I  deemed  myself 
justified  in  doing  so.  Not  that  a  nerve  fibre  proceeds  to  each 
individual  cell  in  the  manner  as  given,  for  several  may  connect 
with  a  nerve  (Fig.  103,  M),  but  that  they  promptly  propagate 
sensory  impressions  produced  by  the  food,  for  setting  up  the 


SERVES   TO   THE   COLUMNAR   EPITHELIUM. 


269 


reflex  actions  in  the  ganglia  for  producing  the  movements  in  the 
muscular  cylinder,  inclusive,  of  course,  of  the  blood-capillaries. 
For  example,  as  soon  as  food  is  introduced  into  the  stomach 


Fig.  103. — A,  B,  C,  D,  E,  isolated  cylindrical  cells  with  processes  containing  nuclei ; 
A,  B,  D,  E,  magnified  590  diameters  ;  C,  magnified  1,200  diameters ;  F,  G,  H, 
cylindrical  cells  with  processes,  which  are  evidently  young  cells,  and  form  atffa 
beautiful  mosaic.    Magnified  1,100  diameters. — Pfluger. 


Fig.  104. — A  Gland  from  the  Submucosa  of  the  Small  Intestine  of  a  Suckling,  ten  days 
old.  a,  Ganglion  ;  b,  nervous  twigs  given  off  by  the  latter  ;  c,  injected  capillary 
network.  This  preparation  had  been  macerated  for  a  very  long  period  in  pyrolig- 
neous  acid. — Frey. 

or  mechanical  irritation  applied  to  the  mucous  membrane — 
e.  g.,  touching  with  a  glass  rod — the  capillaries  expand  and 
the  movements  in  the  walls  begin,  which  would  seem  to  show 
intimate  nervous  connections  subsisting  between  the  epithelium. 


270  NERVES   TO   THE  INTESTINES. 

and  the  ganglionic  layers,  the  same  applying,  of  course,  for 
the  whole  of  the  alimentary  canal,  while  the  investigations 
of  Pfluger  would  show  a  reason  for  it.  The  following  illus- 
tration (Fig.  104)  exhibits  the  appearance  of  a  ganglion  (a) 
from  the  submucosa  or  Meissner's  layer,  with  the  nerves 
radiating  from  it  (&).  The  following  illustration  (Fig.  105)  is 
a  section  of  the  ganglionic  layer  between  the  circular  and  longi- 
tudinal muscles,  by  Auerbach,  and  named  after  him.  Fig.  106 
is  a  similar  section  by  Klein,*  in  which  the  nerves  are  differ- 
ently treated  and  more  highly  magnified. 

Then,  again,  looking  to  the  blood  vessels  and  nerves,  with 
reference  to  the  local  actions,  wTe  can  readily  understand  how 
the  vascular  supply  may  be  increased  or  diminished  in  a  given 
viscus,  or  in  a  portion  of  the  same,  without  interfering  with 
neighboring  organs  or  adjacent  parts,  and  so  as  to  limit  the 
action  to  the  special  work  in  hand,  in  the  measure  of  the  re- 
quirements, which  division  in  labor  necessarily  involves,  as  the 
whole  is  founded  in  individualism  or  automatism  in  the  organs, 
as  before  remarked. 

Thus,  in  the  case  of  the  stomach  (Fig.  110),  the  blood 
is  supplied  from  the  coeliac  axis,  which  is  embraced  by 
the  solar  plexus  (Fig.  109) ;  hence  the  sensory  impressions 
in  the  gastric  mucous  membrane  produced  by  the  food  are 
reflected  to  the  appellate  ganglionic  centres  over  the  vessels 
when  these  are  expanded  correspondingly,  the  pressure  in  the 
arterial  system  causing  them  to  be  filled  instantaneously.  And 
any  portion  of  the  stomachal  mucous  surface  may  be  flushed 
in  order  to  increase  the  action  in  the  part,  whether  in  the  car- 
diac or  pyloric  end  of  the  organ,  by  expanding  the  local 
feeder,  which  is  readily  done.  It  is  very  pretty.  The  vessel 
to  the  great  cut  de  sac,  which  gives  off  the  feeder  to  the 
spleen,  the  separate  ones  to  the  mid-regions  or  central  por- 
tions and  pyloric  end  of  the  organ  (Fig.  110)  and  the  hepatic, 
all  springing  out  of  the  coeliac  axis  with  the  great  semilunar 
ganglia  of  the  solar  plexus  at  its  root  for  compelling  response 
to  local  demands,  with  the  separate  ganglia  along  the  indi- 
vidual vessels  for  reporting  the  demands  and  enlarging  the 
lumen  of  the  vessel.     The  great  anastomosis  of  the  vessels 

*  Hand-Book  for  the  Physiological  Laboratory.—  Burdon-Sanderson. 


NEBVES   TO   THE   INTESTINES. 


271 


Fig.  105. 


-From  the  Small  Intestine  of  a  Guinea-Pig. — Auerbach.    a,  Nervous  inter- 
lacement :  6,  ganglia  ;  c,  lymphatic  vessels  ;  d,  lymphatics. 


Fig.  106. — Auerbach's  Plexus  of  Small  Intestine  of  Human  Foetus,  colored  with  gold. 
The  plexus  consists  of  fibrillated  substance,  and  is  made  up  of  trabecular  of  various 
thicknesses,  which  unite  in  large  placoids.  Nucleus-like  elements  (unformed  gan- 
glion cells)  and  ganglion  cells  are  embedded  in  the  plexus,  the  whole  of  which  is 
inclosed  in  a  nucleated  sheath.     (Oc,  2  ;  obj.,  7.) — Klein. 


272 


NEBVJiS   TO   THE  INTESTINES. 
A 


r  $ 

Fig.  107.— Nerves  to  the  Intestines— partly  ideal.  1,  Ganglionic  layer  of  Meissner  ;  3, 
nerves  from  the  same  to  the  villi  and  columnar  epithelium  ;  3,  nerves  connecting  same 
with  the  ganglionic  layer  of  Auerbach  ;  4,  ganglionic  layer  of  Auerbach  ;  5,  5,  5, 
nerves  connecting  the  intestinal  apparatus  with  the  solar  plexus.  A,  A,  villi  ;  B, 
summit  of  a  lymphoid  follicle  ;  d,  muscularis  mucosae  ;  E,  circular  muscles  ;  /,  longi- 
tudinal muscles  ;  g,  peritoneum  ;  v,  v,  arterial  capillaries;  h,  submucosa. 


NERVES   TO   THE   INTESTIXES. 


^73 


along  the  greater  curvature  provides  against  obstruction, 
should  this  occur  in  either  the  hepatic  or  splenic  branches, 
so  that  compensation  is  readily  effected  by  commensurate 
expansion  in  the  other.     In  this  manner,  then,  afflux  of  arte- 


Pig.  108. — Nerves  of  the  Mesentery  (reduced). — Bougery,  etc.  1,  Root  of  superior 
mesenteric  artery  and  nervous  plexus  (a  portion  of  the  transverse  colon  is  excised, 
in  order  to  show  this  circumstance)  ;  2,  superior  mesenteric  plexus  ;  5,  5,  continua- 
tion of  same  over  the  walls  of  the  vessels  to  the  intestines  ;  A,  intestines  ;  B,  caecum  ; 
C,  appendix  vermiformis  ;  D,  ascending  colon  ;  E,  transverse  colon  ;  F,  descending 
colon. 


274 


NEliVES   TO   THE   INTESTINES. 


Fig.  109. — Solar  Plexus  (reduced). — Bougery,  etc.  The  letters  of  reference  added.  A, 
great  right  semilunar  ganglion  ;  B,  left  ditto  ;  C,  D,  gaDglia  connecting  right  pneu- 
mogastric  nerve  (2)  and  great  splanchnic  (3)  nerve  with  the  ganglion  of  the  superior 
mesenteric  plexus  (E.  5)  ;  F,  ganglion  connecting  right  pneumogastric  and  great 
splanchnic  nerves  (2.  3)  with  the  renal  (G)  and  aortic  ganglia  (H)  ;  4,  coeliac  axis, 
showing  intimate  blending  of  the  right  pneumogastric  nerve  ;  5,  superior  mesenteric 
artery  and  plexus  ;  6,  7,  8,  lesser  splanchnic  nerve,  terminating  in  renal  plexus  :  9, 
renal  artery  and  plexus  ;  10,  diaphragmatic  plexus  and  artery  ;  11,  spermatic  artery 
and  plexus  ;  12,  tendon  of  small  psoas  muscle  ;  13,  eleventh  rib  ;  15,  eleventh  dorsal 
vertebra  ;  16,  crura  of  diaphragm  ;  17,  ligamentum  arcuatum  internum,  the  fibres 
irregularly  divided. 


ARTERIAL    FEEDERS    TO   THE   STOMACH. 


275 


rial  blood  is  compelled  to  be  in  correspondence  with,  the 
local  actions  in  digestion.  And  looking  from  this  to  the 
small  and  large  intestines,  in  which  the  action  begins  later 
on,   it  is  readily  perceived  that  there  is  extension  of   the 


Pig.  110. — The  Cceliac  Axis  and  its  Branches,  the  Stomach  having  been  raised,  and  the 
Transverse  Mesocolon  Removed. — Gray. 

same  principle  in  mechanics  to  them  also.  For  example, 
the  small  intestines  are  supplied  by  the  superior  mesenteric 
artery  (Pig.  Ill),  which  functions  as  the  feeder  to  these  por- 
tions, the  vessel  giving  off  lateral  branches  to  feed  the  vari- 
ous portions,  the  vessels  being  given  off  at  regular  intervals, 


276 


ARTERIAL   Fl.EDERS   TO   SMALL    INTESTINES. 


commencing  with  the  duodenum.  The  transverse  and  ascend- 
ing portions  of  the  colon  and  ccecum  are  supplied  by  the 
colica  media  (11),  colica  dextra  (14),  and  ilio-colica  (15),  given 
off  from  the  opposite  side  of  the  vessel ;  while  the  descending 
colon  and  rectum  are  supplied  by  the  inferior  mesenteric 
artery  (Fig.  112,  9),  the  superior  hemorrhoidal  ( 1 3)  descending 
as  low  as  the  middle  of  the  sacrum,  where  it  divides  into  two 


Fig.  111. — Course  and  Distribution  of  the  Superior  Mesenteric  Artery. — Wilson  and  Bu- 
chanan. 1,  Descending  portion  of  the  duodenum;  2,  transverse  portion;  3,  pancreas; 
4,  jejunum ;  5,  ileum  ;  6,  caecum  and  appendix  vermif ormis ;  7,  ascending  colon  ;  8, 
transverse  colon  ;  9,  descending  colon  ;  10,  superior  mesenteric  artery  ;  11,  colica 
media  ;  19,  the  branch  which  inosculates  with  the  colica  sinistra  ;  13,  pancreatico- 
duodenalis  inferior  ;  14,  colica  dextra  ;  15,  ileo-colica  ;  16,  16,  vasa  intestini  tenuis. 

branches,  which  continue  along  the  sides  of  the  rectum,  divid- 
ing up  into  still  smaller  branches,  to  be  distributed  between  the 
mucous  membrane  and  the  muscles,  nearly  to  the  anus,  anas- 
tomosing with  the  middle  and  inferior  hemorrhoidal  branches 
of  the  internal  iliac  and  internal  pudic  arteries.  By  means  of 
this  arrangement,  then,  together  with  the  local  ganglia  and 
nerves  for  operating  them,  the  vessels  are  readily  expanded 
and  contracted  for  regulating  the  blood-supply.     One  other 


ARTERIAL   FEEDERS   TO    LAHGE   INTESTINES. 


277 


thing !  Bear  in  mind  that  every  nervous  ganglion  is  a  dilator 
and  contractor  centre  for  the  vessels  ;  hence,  it  is  readily  per- 
ceived that  the  actions  can  be  carried  on  in  the  several  portions 
independently,  and  without  involving  other  portions.  For 
example,  expansion  in  the  lateral  branches  to  the  ascending  and 


Pig.  112. — Branches  of  the  Inferior  Mesenteric  Artery. — Wilson  and  Buchanan.  1,  1, 
The  superior  mesenteric,  and  small  intestines  turned  over  to  the  right  side  ;  2,  caecum 
and  appendix  caeci  ;  3,  ascending  colon  ;  4,  transverse  colon  raised  upwards  ; 
5,  descending  colon ,  6,  sigmoid  flexure  ;  7,  rectum  ;  8,  aorta  ;  9,  inferior  mesenteric 
artery  ;  10,  colica  sinistra,  inosculating  with,  11,  colica  media  ;  12,  12,  sigmoid 
branches  ;  13,  superior  hemorrhoidal  artery  ;  14,  pancreas  ;  15,  descending  portion 
of  the  duodenum. 

transverse  colon  (Fig.  Ill,  12,  14,  15)  should  not  affect  the  vas- 
cular supply  to  the  small  intestines,  the  vessels  remaining  at  a 
given  calibre,  since  pressure  in  the  arterial  system  would 
compel  them  all  to  be  filled  simultaneously  But,  since 
the  actions  alternate,  it  is  easy  to  perceive  how  one  set  of 
vessels  may  contract  their  calibre  as  others  expand;   which 


278  VASO-DILA'IOR   AND   CONTKACTOR  NERVES. 

must  be  done  in  order  to  obviate  too  great  depletion  of  the 
arterial  system ;  otherwise  inevitable.  So  that  this  also  has 
its  metes  and  bounds  for  maintaining  a  balance  in  the  organ- 
ism. During  full  digestion  a  vast  amount  of  arterial  blood  is 
thus  poured  into  the  organs  ;  so  that  the  arterial  system  is  com- 
pelled to  condense  itself  in  order  to  maintain  pressure,  tending 
to  produce  brain-ansemia  ;  hence  the  heavy  sleep  of  the  glutton, 
which  is  especially  well  marked  in  cases  in  which  a  degree 
of  anaemia  exists — e.  g.,  convalescence  from  essential  fevers  and 
wasting  diseases.  For  this  reason,  also,  it  acts  beneficially  in 
cases  of  cerebral  irritation  by  calling  off  the  blood,  thereby 
inducing  sleep.  In  the  lower  animals,  in  which  arterial  pres- 
sure is  low,  it  induces  a  semi-comatose  condition  for  many 
days  together,  since  digestion  is  slow  iu  them.  But  in  the 
warm-blooded  animals  provision  is  made  for  digestion  during 
the  maintenance  of  the  activities,  but  which  requires  that 
arterial  pressure  should  be  maintained,  since  it  is  by  means  of 
this  that  the  local  actions  are  increased,  while  <he  necessity  for 
connecting  the  vessels  in  the  intestines  with  the  spinal  medulla 
and  rhythmic  centre  is  to  set  a  limit  to  the  local  actions  and 
maintain  a  balance  in  arterial  pressure  ;  otherwise  the  arterial 
system  itself  might  be  emptied  into  the  viscera,  as  is  seen  in 
"congestive  chill"  and  acute  peritonitis,  in  which  the  action 
is  more  slowly  accomplished,  every  inch  of  ground  contested, 
the  nervous  centres  not  being  suddenly  overwhelmed  as  in  con- 
gestive chill.  The  blending  of  vaso-dilator  and  constrictor 
nerves  in  the  solar  plexus,  and  the  intimate  relations  it  sus- 
tains to  the  cerebro-spinal  axis,  has  its  explanation,  then,  in 
this  circumstance ;  or,  in  other  words,  for  maintaining  a  bal- 
ance in  circulation  during  respiration,  digestion  and  the  vari- 
ous bodily  functions,  the  voluntary  movements  especially; 
while  the  whole  relates  to  evolution  of  force  expended  in  the 
organism. 

M<jde  of  Connecting  the  Double  Ganglionic  Chain  and  Solar 
Plexus  with  the  Central  Nervous  System  or   Cerebro- 
spinal Axis. 
The  following  beautiful  illustration  (Fig.  113)  of  the  fine 
anatomy  in  the  roots  of  the  spinal  nerves  furnishes  a  ready 
explanation  for  the  mode  of  effecting  continuity  in  force  be- 


SPINAL  NERVES  AND   DOESAL   GANGLIA.  27& 

tween  the  viscera  and  central  nervous  system.  The  section 
includes  the  roots  (A,  B)  of  the  spinal  nerves  with  the  gan- 
glion on  the  posterior  root,  carried  beyond  the  point  of  junction 
(C)  of  the  connecting  links  of  the  dorsal  ganglionic  chain., 


Fig.  113.— The  two  Roots  of  the  Spinal  Nerves,  showing  relation  of  the  fibres  of  the 
sympathetic  (C)  to  them.— Ley  dig.  A,  posterior  root ;  B,  anterior  root ;  C,  nerve 
from  the  sympathetic  spinal  chain  at  the  point  of  division  into  the  roots  of  the  spinal 
nerves  ;  trunk  of  a  spinal  nerve  at  the  point  of  union  of  the  two  roots. 

while  the  two  roots  (D)  of  the  spinal  nerves  come  together  to 
form  the  single  nerves  of  which  the  so-called  intercostals  form 
the  major  portion. 

As  will  be  seen,  some  of  the  fibres  of  intercommunication 
(c)  between  the  dorsal  ganglia  and  the   spinal  medulla  pass 


280  SPINAL   NERVES   AND   DORSAL   GANGLIA. 

up  (or  jDass  down,  as  the  case  may  be)  the  anterior  or  motor 
root  of  the  spinal  nerves  (c,  B)}  while  others  pass  up  the  pos- 
terior or  sensory  root  (c,  A),  connecting  with  the  nerve  cells  of 
its  ganglion,  and  through  these  connecting  with  the  spinal  me- 
dulla Thus,  we  have  the  roots  to  the  ganglionic  chain  split- 
ting their  fibrous  bundles  and  dividing  up  into  anterior  and 
posterior  roots,  in  correspondence  with  the  spinal  nerves  with 
which  they  are  blended.  Now,  then,  we  may  readily  under- 
stand how  nervous  force  may  travel  up  and  down  the  gan- 
glionic chain  to  and  from  the  central  nervous  system  readily 
enough,  and  why  sensory  impressions  in  the  mucous  surface 
should  be  promptly  reflected  thence  to  the  cerebro-spinal  axis  ; 
also,  why  sensory  impressions  in  the  skin  should  be  similarly 
reflected,  and  why  stimulation  of  the  cutaneous  surface — e.  g., 
cold  and  hot  applications,  cups,  blisters,  etc. — should  be  bene- 
ficial in  internal  inflammations — pneumonia,  for  example — 
the  impressions  being  reflected  by  the  special  spinal  nerves 
(Fig.  114,  A,  A)  to  the  spinal  medulla,  thence  through  the  in- 
tercommunicating nerves  (S,  S)  to  the  posterior  pulmonic 
plexus  ( P,  P)  to  the  lungs,  producing  contraction  in  the  dilated 
and  engorged  vessels  ;  the  same  remark  applying  for  the  cuta- 
neous surface  and  viscera  in  the  abdomen  as  well. 

The  rapid  condensation  of  the  relaxed  and  bleeding  womb 
from  cold  applications  made  to  the  hypogaster  has  undoubt- 
edly its  explanation  in  this  circumstance,  all  of  which  is  plain 
enough.  But  the  principle  involved  in  this  intimate  nervous 
connection  between  the  internal  and  external  parts  is  the  main- 
tenance of  a  balance  in  circulation  during  the  multiplied  and 
diversified  experiences,  the  cardinal  circumstance  being  to 
bring  it  in  correspondence  with  the  functions  in  the  lungs  and 
digestive  processes,  for  producing  force  in  the  organism  and 
maintaining  a  balance.  The  expeditious  manner  in  which  the 
blood  is  shifted  from  part  to  part  commensurate  with  the  physi- 
ological requirements  in  the  organs  and  organism,  has  its  ex- 
planation in  the  high  pressure  in  the  arterial  system  and  the 
power  of  expanding  the  local  vessels,  the  blood  in  consequence 
rushing  into  them  instantaneously  to  equalize  pressure.  The 
necessity  for  a  vaso-motor  centre  to  effect  expansion  and  con- 
traction in  the  vascular  system  for  coordinating  it  with  the 


SPINAL   NERVES   AND   DORSAL   GANGLIA. 


281 


functions  in  the  lungs,  and  for  maintaining  a  balance  in  circu- 
lation, is,  therefore,  sufficiently  obvious  ;  while  to  expand  and 
contract  the  lumen  inheres  in  the  vessels  themselves,  the  nerv- 
ous apparatus  serving  to  energize  the  actions  and  coordinate 
them  in  the  functions  ;  otherwise,  it  were  utterly  impossible  to 
carry  on  the  functions  in  the  organs. 


Fig.  114.— Transverse  Section  of  Spinal  Cord  and  the  Ganglionic  Chain  of  Sympathetic 
Nerves,  showing  mode  of  connection  between  them.  A,  four  upper  intercostal 
nerves  ;  1,  ganglion  on  posterior  root ;  S,  nerves  connecting  the  ganglia  of  the  sym- 
pathetic with  the  two  roots  of  the  spinal  nerves  ;  P  P,  posterior  pulmonic  plexus  ; 
5,  inferior  cervical  ganglion  ;  C  P,  branches  to  the  cardiac  plexus. 

In  fine,  this  law  of  pressure  and  the  power  of  producing 
rapid  rhythmical  expansions  and  contractions  in  the  vessels 
for  changing  pressure  inheres  in  the  vessels  themselves  to  the 
minutest  capillaries,  and  by  means  of  which  circulation  is  made 
to  respond  to  the  physiological  requirements  in  the  organs  and 
organism,  otherwise  impossible ;  while  for  producing  equilib- 


282  YASO-DILATOR   AND   CONTRACTOR   NERVES. 

rium  it  requires  the  correlation  of  the  two  nervous  forces 
which  these  opposite  actions  represent  in  the  medulla  oblon- 
gata— indeed,  in  every  nervous  ganglia  or  centre  of  the  local 
actions. 

The  point' we  wish  to  make  is,  that  while  opposite  nervous 
forces  are  thus  indicated  in  the  two  roots,  the  ganglion  itself 
possesses  independent  action  for  producing  the  opposite 
movements  in  the  muscles,  otherwise  inexplicable  ;  proving 
conclusively  a  correlation  of  the  forces  in  the  ganglion  and  the 
power  of  coordinating  the  local  actions,  while  this  again  is 
augmented  by  the  force  through  the  roots  for  energizing  the 
movements.  And  it  is  a  notable  circumstance  that  the 
spinal  ganglia  possess  two  roots  connecting  them  with  the 
spinal  nerves  (Fig.  116).  Not  that  they  necessarily  are  op- 
posing lines  of  force,  for  each  root  may  contain  both  sets  of 
fibres  ;  but  the  fact,  nevertheless,  is  deeply  suggestive. 

Accordingly,  I  have  ventured  to  connect  the  columnar 
epithelium  in  the  intestines  with  the  nerves  proceeding  from 
the  ganglionic  layer  of  Meissner  (Pig.  107,  1,  2).  This  por- 
tion of  the  picture  is,  therefore,  ideal.  But,  considering  the 
rapidity  of  secretory  and  absorptive  functions  in  the  intes- 
tines, the  quick  response  to  stimuli,  I  have  felt  justified 
in  extending  the  nerves  to  the  muscles  and  capillaries  (of 
which  there  can  be  no  question)  to  the  adjacent  epithelium. 
Not  that  it  occurs  in  the  exact  manner  here  depicted,  but 
that  something  similar  to  this  does  actually  exist,  I  hold  to  be 
a  reasonable  deduction  from  the  evidence  in  the  salivary 
glands.  Still,  it  is  not  essential  to  the  argument,  for  we  know 
the  cells  have  power  to  expand  and  contract  in  the  entire  ab- 
sence of  muscles  and  nerves  for  effecting  it  ;  only  it  would 
seem  to  be  necessary  for  bringing  them  into  correspondence 
and  for  increasing  their  functions.  And  to  bring  the  muscular 
walls  and  the  blood  vessels  into  correspondence  with  the  ac- 
tions in  the  mucous  membrane,  is  the  explanation  for  the  inter- 
communicating nerves  between  the  ganglionic  layers  of  Meiss- 
ner and  Auerbach  (3),  and  between  the  latter  and  the  solar 
plexus  (Figs.  108,  5,  5,  5  ;  109,  5,  5,  2,  1).  The  lims  of  nerv- 
ous force  to  and  from  the  solar  plexus  are  upon  the  blood- 
vessels ;  hence,  it  it  is  easy  to  perceive  how  these  are  brought 


NEKVES   TO   VESSELS   AND   TJSCEEA   IN  ABDOMEN.         283 

into  correspondence  with  the  physiological  requirements  in  the 
organs,  or  with  the  digestive  and  absorptive  processes.  Hence, 
as  soon  as  food  is  introduced  in  the  stomach,  or  mechanical 
irritation  is  applied  to  the  mucous  surface — e.  g  ,  touching 
it  with  a  glass  rod — the  capillaries  expand  and  the  movements 
in  the  muscular  walls  begin. 

The  intimate  relations  which  the  intestines  sustain  to  the 
cerebro-spinal  axis  and  respiration  may  be  seen  in  the  blend- 
ing of  the  pneumogastric  and  splanchnic  nerves  in  the  solar 
plexus  (Fig.  ]  09,  2,  3,  b). 

Finally,  it  is  well  known  that  stimulation  of  the  pneumo- 
gastric nerves  produces  peristalsis,  the  same  remark  applying 
to  the  ganglia  in  the  solar  plexus,  and  that  stimulation  of  the 
pneumogastric  nerves  produces  expansion,  while  stimulation 
of  the  splanchnic  nerves  produces  contraction  in  the  vascular 
network  in  the  mucous  membrane  of  the  stomach  and  intes- 
tines, showing  conclusively  the  power  in  the  medulla  oblon- 
gata to  produce  the  movements  in  the  intestines,  and  to  ex- 
pand and  contract  the  vessels  in  the  mucous  membrane,  the 
same  as  in  the  cutaneous  surface  and  systemic  vessels  ;  other- 
wise, it  were  utterly  impossible  to  maintain  a  balance  in  the 
circulation,  or  carry  on  the  functions  in  the  organs,  the  one 
involving  the  other. 

"Inhibitor  centre!"  -'Inhibitor  nerves!"  Inhibit  what? 
The  heart  expands  and  contracts  in  conformity  with  the  law 
for  changing  pressure.  In  which  direction,  then,  does  your 
"inhibitor"  act?  You  irritate  the  pneumogastric  branch, 
and  because  diastole  is  made  excessive  thereby,  producing  an 
intermittent  pulse,  you  term  it  an  inhibitor  nerve,  whereas  the 
heart  is  expanded  to  its  utmost  limit,  when  the  "  cardiac  in- 
hibitor" is  stimulated,  and  when  excessive  producing  death, 
the  heart  in  extreme  diastole,  while  the  response  is  prompt 
and  energetic  ;  moreover  it  performs  work,  reducing  pressure 
in  the  organ,  the  blood  in  consequence  rushing  into  it  and  fill- 
ing it  instantaneously.  And  yet  it  is  to  count  for  nothing.  IS'  a- 
ture  withdraw  force  from  the  heart  and  blood-vessels,  let  go, 
as  it  were,  yet  controlling  the  blood  and  compelling  circulation 
in  the  measure  of  the  physiological  requirements  in  the  lungs, 
in  the  heart,  in  the  blood-vessels,  in  the  cell-brood,  and  in  the 


284 


VASO-DILATOR   NERVES   DEMONSTRATED. 


very  blood  itself,  as  has  been  fully  set  forth  !  A  pretty  let  go 
that  is.  Fie  on  it !  Nature  works  by  law  ;  your  theory  does 
not,  and  you  must  bring  it  into  correspondence,  else  stay  in 
outer  darkness.  You  offer  no  reason  for  it,  whereas  there  is  a 
reason  for  everything  that  Nature  does.  It  is  a  true  saying : 
"  One  lie  is  the  father  of  many."  But  Truth  is  a  sleuth-hound 
— terrible,  and  the  progeny  is  doomed. 

This  action  of  the  nerves  upon  the  vessels  has  forcible  illus- 
tration in  the  case  of  the  vessels  and  nerves  to  the  salivary 
glands,  notably  the  submaxillary  and  sublingual  glands  (Fig. 
115).     Thus,   when  the  chorda  tympani  (c)  is  irritated,   the 

T 


Fig.  115. — Nerves  of  the  Submaxillary  and  Sublingual  Glands  of  the  Dog. — Bernard. 
N,  submaxillary  gland  ;  O,  sublingual  gland  ;  JM,  Wharton's  duct,  in  which  a  can- 
ula  has  been  placed  ;  JL,  duct  of  the  sublingual  gland,  also  furnished  with  a 
canula  ;  1\  S,  S',  the  lingual  branch  of  the  fifth  nerve  ;  F,  the  facial  nerve  ;  c, 
chorda  tympani ;  g,  the  submaxillary  ganglion  ;  q,  the  superior  cervical  ganglion  ; 
P,  sympathetic  twig  passing  from  the  ganglion  to  the  submaxillary  gland  ;  j,  inter 
nal  maxillary  artery  ;  V,  vidian  nerve  ;  I,  branch  of  the  lingual  nerve  ramifying  in 
the  buccal  mucous  membrane. 

arteries  of  the  gland  dilate,  the  blood  surging  into  them 
under  the  pressure  in  the  arterial  system,  while  the  veins  lead- 
ing from  the  organ  are  made  to  pulsate  under  the  force,  and 
when  they  are  divided  the  blood  spurts  like  in  an  artery.  But 
when  the  sympathetic  fibres  from  the  cervical  ganglion  (q,  P) 
are  excited,  the  arteries  contract  and  circulation  in  the  gland 
is  retarded  correspondingly  ;  and  if  the  veins  are  now  cut  they 
discharge  "black"  blood  in  a  slow  stream. 


VASO-DILATuR  NEKVES   DEMONSTBATED.  285 

JN'ow,  then,  when  the  chorda  is  irritated  force  is  not  with- 
drawn from  the  blood-vessels  any  more  than  in  the  cervical 
fibres  when  they  are  irritated,  only  that  different  7c  inds  of  elec- 
trical force  pass  along  the  nerves,  positive  or  negative,  as  the 
case  may  be,  but  opposite  kinds  of  force  evidently  for  produc- 
ing the  opposite  movements  in  the  vessels,  according  to  whether 
it  is  desirable  to  increase  or  to  diminish  the  circulation  in  the 
parts,  and  which,  of  course,  is  determined  by  the  sensory  impres- 
sions propagated  from  the  mucous  membrane  of  the  mouth  by 
the  food  for  increasing  the  saliva  during  mastication,  the  flow  di- 
minishing with  this,  and  ceasing  nearly  altogether  when  there 
is  no  stimulus  in  the  mouth  for  producing  sensory  impressions. 
Furthermore,  there  must  be  prompt  response,  for  food  cannot 
be  kept  forever  in  the  mouth,  and  the  saliva  is  needed  for  pre- 
paring the  bolus  and  making  dry  substances  moist  so  they  can 
be  swallowed. 

Finally  the  ganglia  or  mind-centres  have  both  sets  of  fibres 
— vaso-dilator  as  well  as  vaso-constrictor  nerves — running  into 
them,  which  is  essential  for  effecting  coordination  and  main- 
taining a  balance  in  the  local  circulation ;  otherwise  impos- 
sible. For  example,  the  submaxillary  ganglion  (I^ig.  115,  g) 
connects  with,  the  chorda  tympani,  c'  (a  branch  of  the  facial), 
and  lingual,  T,  8,  S'  (a  branch  of  the  fifth  pr.),  upon  the  one 
hand  and  with  the  superior  cervical  ganglion  (q,  P,  g)  upon 
the  other.     All  which  is  plain  enough. 

You  are  over  anxious  about  contraction,,  but  seemingly  in- 
different about  expansion  ;  and  yet  the  fact  is  undeniable  that 
without  the  latter  you  could  nut  have  the  former — nay,  more 
than  this,  that  it  must  have  precedence  or  first  place,  whether 
it  relate  to  fluids  or  solids,  living  or  non-living  matter.  Who 
denies  that  his  opinion  is  not  entitled  to  respect,  for  he  talks 
without  authority  and  without  knowledge.  Being  correlated 
forces  in  Nature,  they  are,  therefore,  inseparable  ;  while  in  the 
animal  organism  the  tissues  are  kept  in  constant  motion  by 
their  alternating  actions,  whether  it  relate  to  circulation  or  the 
voluntary  movements  while  coordination  is  effected  by  means 
of  the  appellate  ganglionic  nervous  centres  for  the  organs  and 
organism. 

So,  then,  the  power  for  coordinating  the  viscera,  inclusive  of 


286  THE   PNEUMOGASTRIC   AND   SPLANCHNIC   NERVES. 

the  blood-vessels,  inheres  in  the  medulla  oblongata,  the  rhyth- 
mic centre  for  the  organism. 

The  following  instructive  diagram  (Fig.  116)  will  give  some 
idea  of  the  vast  number  of  the  sympathetic  nerves  and  gan- 
glia to  the  vessels  and  viscera  in  the  abdomen.  The  spinal 
ganglionic  chains  (one  side  (left)  only  rei>resented  in  the 
figure)  connect  with  the  solar  plexus  by  means  of  the 
splanchnic  nerves  (extended  links  indicating  where  the  gang- 
lia belong  in  the  chain  in  the  respective  sides)  ;  below,  they 
connect  with  the  aorta  and  lower  cava,  following  the  branches 
into  the  viscera,  increasing  in  numbers  as  we  approach  the 
pelvic  viscera,  the  occasion  for  which  will  appear  later  on ; 
above,  with  the  viscera  and  vessels  in  the  chest,  and  blending 
with  the  pneumogastric  and  phrenic  nerves  in  the  solar  plexus. 
In  this  manner,  then,  the  whole  is  consolidated,  and  the  action 
in  the  viscera  is  unified  with  the  central  nervous  system,  so 
that  a  balance  is  readily  maintained  in  the  circulation  during 
respiration  and  the  voluntary  movements,  etc. 

In  other  words,  the  parts  are  unified  through  the  nervous 
apparatus.  The  question  as  to  which  of  the  two  lines  of  nerv- 
ous force  from  the  central  nervous  system  to  the  stomach  and 
intestines — this  by  way  of  the  pneumogastric  nerves  or  that  of 
the  splanchnics — is  the  more  important,  would  seem  to  be  in 
favor  of  the  former,  judging  from  the  evidence  before  us  and  our 
own  observations  ;  though,  perhaps,  a  true  comparison  cannot 
be  made  for  the  reason  that  the  two  sets  of  fibres,  or  dilators  and 
contractors,  are  not  equally  distributed  between  them,  the 
pneumogastrics  containing  more  of  the  former,  the  splanchnics 
more  of  the  latter  fibres ;  while  the  one  relates  more  particularly 
to  the  action  in  the  muscular  cylinder,  the  other  the  vascular 
apparatus  ;  while  neither  are  exclusive  in  their  functions.  But 
strong  currents  of  nervous  force  set  in  and  out  of  the  viscera 
to  and  from  the  central  nervous  system  through  the  pneumo- 
gastric nerves,  for  producing  correspondence  between  them 
and  the  containing  walls,  at  the  same  time  increasing  the 
actions  in  the  muscular  cylinder  and  vascular  apparatus,  ex- 
panding the  network  of  capillaries  and  producing  peristalsis  ; 
since  stimulation  of  the  pneumogastric  nerves  produces  these 
effects,  while  stimnlntion  of  the  splanchnic  nerves  produces 


ACTION   IN   'IHE   J'NJiUMOGASTKlCS.  237 

contraction  of  the  capillary  network,  though  it  also  produces 
peristalsis ;  but  they  cannot  effect  the  actions  in  the  stomach 
nor  set  up  the  movements  in  the  medulla  oblongata  during 
ingestion  for  bringing  the  containing  walls  into  correspondence 
with  the  exrjansile  action  in  the  siomach  for  producing  the 
requisite  room  for  the  food  and  maintaining  a  balance  in 
pressure,  otherwise  impossible ;  at  the  same  time  expanding 
the  capillary  network  in  the  gastric  mucous  membrane  for 
increasing  the  digestive  and  absorptive  processes  in  the  organ. 
It  comes  to  this  in  the  main :  that  the  pneumogastric  nerves 
produce  expansile  action  in  the  viscera,  while  the  splanchnics 
have  the  opposite  effect— producing  contraction,  especially  in 
the  vessels.  And  when  the  pneumogastrics  are  divided  in  the 
neck,  the  stomach  and  abdomen  can  no  longer  be  expanded 
nor  any  food  be  taken,  the  animals  dying  from  thirst  and 
starvation,  precipitated  by  pulmonary  congestions  from  two  to 
five  days,  the  nerves  not  uniting,  which  has  occasionally  hap- 
pened ;  while  the  respiratory  rhythms  fall  from  twenty-five  to 
thirty  per  minute  in  the  dog  to  seven  and  eight,  sometimes 
as  low  as  two  per  minute.  But  important  corroborative  evi- 
dence is  obtained  also  at  the  bedside  in  cases  of  spinal  injury, 
in  which  the  nervous  currents  through  the  spinal  cord  are  cut  off, 
thus  throwing  the  burden  of  effecting  coordination  in  the  stom- 
ach upon  the  pneumogastrics,  and  showing  the  great  role  they 
perform  in  the  mechanics  in  the  abdomen,  as  well  as  in  the  chest, 
in  connection  with  respiration  and  circulation,  which  has 
already  been  fully  considered.  Thus,  in  a  case  of  spinal  injury 
which  came  within  my  own  knowledge,  in  which  the  fourth 
and  fifth  cervical  vertebrae  were  fractured  and  displaced  so 
as  to  crush  the  cord,  producing  complete  paralysis,  with 
anaesthesia  from  the  clavicles  down,  a  little  sensation  only  being 
perceptible  about  the  clavicles,  not  knowing,  indeed,  "which 
portion  of  the  body  was  in  contact  with  the  bed  but  by  actual 
sight,"  as  he  said,  commenting  upon  it,  and  deeming  it  very 
curious,  the  patient  living,  however,  over  twenty  days,  the 
following  facts  were  ascertained  ;  notably  :  1 .  Respiration  was 
diaphragmatic  and  labored,  shallow,  and  from  25  to  30  per 
minute  ;  pulse,  120  to  130  per  minute ;  polyuria,  with  complete 
paralysis  of  the  bladder,  the  urine  having  to  be  drawn  every 


283         NERVES    TO   VESSELS   AND   VISCEBA   IN   ABDOMEN. 


/hur  T.'DtrjalNenit 
Tha  Z'.'E'iinlAttur 
/ToatSf'IltrinlM 


pom  I'/Sntm/Mnir 

Jl-Mm  Z7*Saertithh-ur 

rnnrt^Sarml  Af/rv, 

fres>S$£nriH. 

It-em  (oectjfftft  Yrm 


Oii^glnvftAaGinUiAiTehtU 


Fig.  116.— The  Abdominal  Sympathetic  Nerves  in  Diagram.— Flower. 


EXTENSIVE   DISTRIBUTION   OF  THE   PNEUMOGASTRICS.     239 

three  or  four  hours,  the  patient  being  made  aware  of  the  pass- 
ing of  the  instrument  by  sight  only ;  tendency  to  diarrhoea, 
but  no  sensation  about  the  anus,  the  patient  cognizant,  how- 
ever, when  an  action  of  the  bowels  was  expected,  directing 
the  attendant  to  the  circumstance ;  and  when  the  bed-pan  was 
placed  under  him,  presently  there  would  be  an  action.  Occa- 
sionally he  relished  food ;  there  was  considerable  venous  stasis  in 
the  systemic  capillaries,  with  tendency  to  serous  effusions  in  the 
connective  tissue,  the  result  rather  of  defective  absorption.  He 
lingered  twenty-one  days,  and  the  post-mortem  examination 
showed  the  nature  of  the  injury  to  the  spinal  cord,  which  was 
crushed  and  completely  disorganized  at  the  seat  of  injury. 
JN'ow,  then,  attention  is  specially  directed  to  the  interesting 
oircumstances  italicized  in  the  notes — namely,  the  polyuria 
and  the  sensory  impressions  transmitted  from  the  colon  during 
a  faecal  action. 

Of  course,  the  polyuria,  with  the  tendency  to  diarrhoea,  should 
be  explicable,  since  they  flow  out  of  the  mechanics  as  a  result  of 
the  lesion  in  the  spinal  cord,  and  are  easily  explained.  The 
spinal  cord  being  crushed,  this  cuts  off  the  nervous  force  from 
above  through  the  splanchnic  nerves,  so  that  the  cervical  chain 
and  cardiac  plexus  must  be  the  route  of  the  nervous  supply  ; 
this,  together  with  the  contractor  fibres  in  the  pneumogastric 
nerves,  which  is  not  sufficient  for  maintaining  the  lumen  and 
tonus  in  the  vessels  ;  in  consequence,  an  amount  of  dilatation 
results,  producing  unusual  fullness  in  the  vessels  and  hy- 
peremia in  the  organs ;  hence,  the  polyuria  and  the  tendency 
to  diarrhoea.  Then,  again,  the  embarrassed  respiration  also 
t^nds  to  produce  fullness  in  the  portal  vessels  and  lower  cava 
system,  which  should  increase  the  secretory  actions,  so  that  it 
is  not  at  all  difficult  to  account  for  the  polyuria  and  tendency 
to  diarrhoea,  under  the  circumstances.  The  consciousness  of 
the  faecal  action  is  undoubtedly  produced  through  the  pneu- 
mogastric nerves.  The  contiguity  of  the  transverse  colon  to 
the  stomach  suggests  direct  extension  of  the  nerves  to  the 
colon,  the  left  pneumogastric  nerve  especially,  which  is  dis- 
tributed principally  over  the  anterior  poriions  of  the  stomach  ; 
but  in  order  to  do  this,  the  nerves  would  have  to  penetrate 
both  layers  of  peritoneum,  gastric  and  colonic  ;   besides,  it 


290     EXTENSIVE    DISTRIJHTION    OF   THK   PNEUMOGASTRICS. 

'  would  interfere  with  the  free  action  of  the  organs.  The  nerv- 
ous force,  then,  passes  through  the  solar  plexus,  thence  up 
the  colonic  ligament,  colica-media,  colica-dextra,  colica-sini  tra, 
and  inferior  mesenteric  arteries  to  the  colon  ;  showing,  also, 
that  the  current  is  not  broken  at  the  ganglia,  but  passes  to  and 
fro  between  the  viscera  and  central  nervous  system  without  let 
or  hindrance.  The  pain  in  colic  from  over-distension  of  the 
colon  by  the  gases,  and  the  traction  upon  the  nervous  filaments 
which  this  produces,  has  similar  explanation  ;  the  sensory  im- 
pressions being  propagated  over  the  pneumogastric  nerves,  and 
so  promptly  reaching  the  brain  And  the  lungs  become  filled 
up  and  carnified  after  division  of  the  pneumogastric  nerves, 
for  the  reason  that  the  fibres  from  the  posterior  pulmonic  and 
cardiac  plexuses  are  not  sufficient  for  maintaining  the  normal 
lumen  in  the  capillary  network  in  the  alveoli  with  the  pneu- 
mogastric fibres  divided,  the  vessels  consequently  become  en- 
gorged, leading  to  effusions  and  haemorrhage  ;  hence  this  cir- 
cumstance. According  to  Latschenberger  and  Deahna,*  the 
vagus  (pneumogastric)  contains  both  pressor  and  depressor 
(vaso-contractor  and  vaso-dilator)  fibres ;  consequently,  we 
should  have  congestions  in  the  vessels  of  the  chest  and  abdo- 
men after  division  in  the  neck  ;  and  the  pulmonic  congestions 
should  therefore  have  this  explanation  :  the  flood-gates  thrown 
open,  we  must  have  inundations  in  the  lungs  as  well  as  in  the 
abdomen  flowing  out  of  this  circumstance,  to  the  contrary, 
notwithstanding.  At  any  rate,  it  would  explain  the  pulmonic 
congestions  and  the  other  wrould  not,  which  places  it  under  a 
cloud.  The  thick  bundles  of  nerve  fibres,  continued  from 
the  pneumogastric  nerves  to  the  central  ganglia  of  the  solar 
plexus  (Fig.  109,  2,  (7,  D,  E),  are  reflected  thence  over  the 
cceliac  axis  (4,  4)  to  the  stomach,  liver,  spleen  and  pancreas  ; 
and  over  the  mesenteric  artery  (5)  to  the  small  and  large  in- 
testines, the  cffical  and  sigmoid  portions,  by  way  of  the  inferior 
mesenteric  artery  ;  while  that  by  the  splanchnic  nerves  (3,  H) 
passes  by  way  of  the  semilunar  (A,  B)  and  central  ganglia 

Concerning  the  Action  in  the  Kidneys. — Now,  then,  let  us 

*  Latschenberger  u.  Deahna,  PfliXgefs  Arehiv,  vol.  xiii.,  p.  22. 


PHYSIOLOGICAL   ANATOMY    OF   THE   KIDNEYS.  291 

look  from  the  mechanics  in  the  intestines  to  the  action  in  the 
kidneys  for  interpreting  the  special  phenomena,  anatomical 
and  physiological,  appertaining  to  them  also.  Briefly,  the 
renal  artery  (which  is  a  very  large  vessel),  previous  to  entering 
the  kidney,  divides  np  into  four  or  five  branches  that  pass  into 
the  organ  at  the  hilus — the  vein  in  front,  the  ureter  behind  ; 
and  after  dividing  and  subdividing  to  form  the  arterioles 
rectce  between  the  tubili  recti,  send  off  lateral  branches  (Fig. 
117,  ai,  va),  which  terminate  in  the  capillary  tufts  inclosed  in 
the  expanded  ends  of  the  convoluted  tubes  known  as  corpora 
Malpighiana  {gl)  ;  and  from  which  proceeds  an  efferent  vessel 
(?>c),  which  also  breaks  up  into  a  capillary  network  around  the 
convoluted  tube  (c),  and  medullary  substance,  or  very  much  as 
the  portal  vein  in  the  liver,  and  winch  again  converge  in  a  com- 
mon vein  (v,  z)  that  discharges  into  the  interlobular  vein, 
or  vena  recta,  side  by  side  with  the  arteriola  recta  (m). 

The  convoluted  tubes  are  composed  of  two  layers,  the  mem- 
br ana  propria  and  an  epithelium,  consisting  of  a  single  layer 
of  polygonal  nucleated  cells,  in  which  metamorphosis  is 
effected.  The  epithelium  occupies  about  two-thirds  the  diam- 
eter of  the  tube,  the  remainder  representing  the  lumen  ;  but  that 
within  the  glomerulus  itself  is  thinner.  Finally,  these  convo- 
luted tubes  lying  in  the  cortical  portion  are  gathered  up  into 
the  straight  bundles  that  form  the  conical  masses  known  as 
the  pyramids  of  Malpighi  (Fig.  118,  3),  coalescing  and  joining 
at  acute  angles  at  the  bases  and  within  the  pyramids  to  bring 
about  this  result,  each  one  containing  from  two  to  five  hundred 
tubes,  which  open  upon  the  apices  (4)  of  the  pyramids  (from 
eight  to  fifteen  in  number),  that  project  into  the  calices  and 
pelvis  of  the  kidney. 

Now,  then,  the  effect  of  this  arrangement  is,  when  the  lumen 
of  the  renal  artery  is  enlarged  for  increasing  the  action  in  the 
organ,  it  first  flushes  the  capillary  tufts  in  the  glomeruli 
(Fig.  117,  va,  gl),  extending  thence  to  the  vascular  plexuses  of 
the  convoluted  tubes  (ve,  c) ;  but  as  the  tufts  are  the  proximal 
portions,  of  course  arterial  pressure  should  be  more  effective 
here,  tending  to  strain  out  the  substances  that  pass  easily 
through  the  animal  membranes,  passing  out  with  the  stream  of 
water  in  which  they  are  suspended,  or  down  the  tubes,  washing 


292 


PHYSIOLOGICAL   ANATOMY    OF   TIIK    KIDNEYS. 


out  the  secretions  as  it  goes  along;  while  the  remainder,  which 
is  under  reduced  pressure  by  reason  of  this  depletion,  passes 
out  through  the  efferent  vessel  to  the  capillary  network  of 


Fig.  117.— Course  of  the  Blood-Vessels  within  the  Cortex  Proper  (diagrammatic),  m, 
The  space  occupied  by  the  medullary  radius  ;  b,  that  occupied  by  the  convoluted 
canals  ;  ai,  arteria  inter-lobularis  ;  vi,  vena  iuter-lobularis  ;  va,  vas  afferens  glome- 
ruli ;  ve,  vas  efferens  glomeruli ;  gl,  glomerulus  ;  vz,  venous  twig  of  the  inter-lobular 
vein.  — Ludwig. 


Fig.  118.— Section  of  the  Kidney,  surmounted  by  the  Supra-Renal  Capsule.— Wilson.  1, 
Supra -renal  capsule  :  2.  vascular  portion  of  the  kidney  ;  3,  3,  tubular  portion,  con- 
sisting of  cones  ;  4,  4,  papilla  projecting  in  the  caliches  ;  5,  5,  5,  infundibula  ;  6, 
pelvis ;  7,  ureter. 

the  tubes,  the  slower  circulation  in  them  affording  opportu- 
nity for  effecting  metamorphosis  in  the  epithelium,  as  in  the 
case  of  the  portal  circulation,  to  which  it  sustains  close  resem- 


THE   ACTION   IJST   THE    UEETEBS   AND    BLADDER.  293 

hlance  in  more  respects  than  one,  since  the  rhythmical  com- 
pression of  the  liver-substance  during  respiration  for  increas- 
ing the  venous  circulation  in  the  liver,  should  have  the  same 
effect  upon  the  venous  circulation  in  the  kidneys,  for  the 
principle  is  precisely  the  same  The  gentle  and  uniform  com- 
pression of  the  kidneys  during  inspiration  must  certainly 
compel  the  blood  into  the  renal  veins,  while  the  suction  force 
in  the  chest  should  pull  it  toward  the  lungs — the  same  as  the 
portal  blood,  and  the  blood  in  the  hepatic  veins.  Of  course, 
the  same  mechanics  also  applies  for  the  secretions,  tending  to 
force  them  out  into  the  pelvis  of  the  kidney  (Fig.  118,  6).  But 
here  we  have  to  note  a  special  arrangement  that  obtains  for 
increasing  the  action,  constituting  the  fine  adjustment  in  this 
mechanics  ;  notably,  the  action  in  the  pelvis  and  ureter.  The 
ureter,  which  is  expanded  at  the  kidney  to  form  the  pelvis 
and  infundibula  (6,  5),  is  a  muscular  organ  richly  supplied 
with  nerves  from  the  renal  plexus  (Fig.  1(.;9,  9),  while  the 
special  ganglion  or  mind-centre  {G)  directs  the  reflex  actions  ; 
and  there  is  but  little  reason  to  doubt  that  by  means  of  this 
systematic  arrangement  there  is  a  pumping  action  going  on 
in  the  pelvis  of  the  kidney,  much  in  the  same  manner  as  takes 
place  in  the  expanded  buccal  cavity  of  the  leech  during  imbi- 
bition for  aspirating  the  blood  in  the  capillaries,  or  the  still 
more  striking  example  furnished  in  the  nursing  infant  aspi- 
rating the  milk  through  the  lacteal  tubes,  the  papilla  of  the 
mammary  gland  bearing  close  resemblance  to  the  renal 
papillae  which  project  into  the  cavity  of  the  pelvis  for  the 
purpose.  So  that,  while  the  infant  is  aspirating  the  mammary 
gland,  the  ureters  at  the  same  time  are  aspirating  the  kid- 
neys. Is  it  fanciful  %  Answer  this  question,  then :  What  is 
this  mind-centre  doing  here,  and  what  are  muscles  and  nerves 
doing  in  the  ureter,  since  they  cannot  be  purposeless  %  Can 
it  be  doubted  for  a  single  moment,  even,  that  they  relate  to 
the  functions  in  the  kidneys,  designed  to  expedite  them  ?  I 
think  not.  And  wherever  found,  muscles  and  nerves  perform 
.work  in  correspondence  with  the  special  physiological  require- 
ments for  which  they  are  the  relative  adjustments,  here  as 
elsewhere.  What  matters  it  where  they  are  located  ?  Work 
is  their  office,  and  work  they  do.     Then,  too,  look  at  the  shape 


294  THE   ACTION    IN   THE    ETKETER8    AND    BLADDKK. 

of  the  expanded  portion  of  the  ureter  (Fig.  118,  C),  and  com- 
pare the  muscles  with  the  muscles  in  the  leech  and  oesophagus  ; 
notably,  the  internal  layer  of  circular  and  the  external  layer 
of  longitudinal  muscles,  layer  for  layer,  in  them  all,  while  each 
is  operated  by  nerves  from  a  special  ganglionic  nervous  cen- 
tre, for  producing  and  coordinating  the  actions,  or  the  same 
as  the  pneumogastric  nerves  and  ganglia  in  the  case  of  the 
oesophagus,  the  special  arrangements  which  obtain  in  them 
being  adaptive  changes  to  meet  the  requirements  in  the  case. 
Keeping  in  mind,  also,  that  the  kidney,  with  the  afferent  and 
efferent  vessels  and  discharging  duct  is  an  entity,  or  inde- 
pendent organ,  therefore  endowed  with  automatic  action, 
which  is  essential  for  maintaining  the  life  that  is  in  it  and 
carrying  on  its  functions,  otherwise  impossible.  And  it  is 
well  to  look  over  the  organs  in  this  way,  for  pressure  is  the 
basis  of  all  of  them,  and  must  be  so,  in  the  very  nature  of 
things.  Well,  what  does  it  all  mean,  if  not  to  act  alike  in 
pumping  the  fluids  into  themselves  in  connection  with  the 
special  functions,  pressure  applying  to  all  alike  %  Nay,  there 
is  other  evidence  proving  they  mitst  act  alike  ;  otherwise,  it 
would  be  utterly  impossible  to  carry  on  the  functions  in 
the  kidneys,  since  the  uriniferous  tubes  would  be  choked 
from  the  accumulations  in  the  pelvis  from  inability  to 
effect  expansion  in  the  urinary  bladder  at  the  distal  end  of 
the  ureters,  which  must  expand  in  similar  manner  under  the 
stimulus  of  the  urine  for  receiving  it.  Thus  the  waves  of 
expansion  and  contraction,  passing  along  the  ureters  to  the 
bladder,  causes  the  bladder  to  expand,  the  waves  losing  them- 
selves in  the  general  expansion  which  results  in  the  bladder,  or 
the  same  as  in  the  stomach,  when  ingesta  passes  into  it  from 
the  oesophagus,  finally  the  same  as  in  the  worms,  notably  the 
leech  (Fig.  16,  1,  2,  3),  the  structures  being  also  homologous, 
the  principle  the  same  ;  otherwise,  it  would  be  utterly  impos- 
sible to  introduce  the  urine  into  the  bladder.  Besides,  the 
urinary  bladder  is  rich  in  all  the  elements  in  arterial  tissue  ; 
namely,  unstriped  muscles,  elastic  and  connective  tissue  fibres, 
the  bundles  running  in  every  direction,  and  when  contracted,  as 
in  the  empty  condition,  the  structure  is  very  dense.  And  to  think 
of  straining  it  open,  as  when  fully  distended  with  Urine,  by  the 


THE   ACTION   IN   THE   URETERS   AND   BLADDEE.  295 

action  in  the  ureters  and  kidneys,  thereby  producing  strain  to 
the  delicate  tubuli  recti  and  convoluted  tubes  extending  back 
to  the  glomeruli,  or  through  and  through  the  organ  from  hilus 
to  cortex,  is  nothing  short  of  madness.  The  force  is  not  there 
for  expanding  the  bladder,  "but  upon  the  ground,  where  the 
work  is  done  in  the  walls  of  the  viscus  itself,  otherwise  it 
were  utterly  impossible  to  introduce  the  urine  into  the  bladder, 
or  operate  the  kidneys. 

The  strong  muscular  walls  of  the  viscus  are  not  forced  into 
extension  by  the  ureters  and  kidneys.  Aeh  Gott !  Never 
in  tliat  way,  under  the  canopy  of  heaven !  Impossible ! 
Then,  if  you  would  not  have  the  urine  damming  back  upon 
the  delicate  renal  structures,  thereby  arresting  the  functions 
in  these  organs,  the  ureters  in  common  with  the  bladder  must 
respond  to  the  stimulus  of  the  urine ;  the  same  as  the 
oesophagus  and  stomach,  the  cystic  ducts  and  gall-bladder, 
under  the  action  of  the  special  stimulus  in  the  organs,  the 
structures  being  fundamentally  the  same,  and  pressure  apply- 
ing alike  to  all  of  them. 

Then,  again,  as  the  bladder  fills  and  expands  with  the  urine,  it 
rises  into  the  cavity  of  the  abdomen ;  and  when  the  con- 
tents are  being  discharged,  it  sinks  again  to  the  bottom  of 
the  pelvic  basin.  Accordingly,  we  have  the  longitudinal  mus- 
cles increased  in  the  ureters  as  the  bladder  is  approached, 
and  the  addition  of  an  internal  layer,  so  that  the  tubes  may 
readily  elongate  and  contract  under  the  action  of  the  special 
nerves  in  the  parts,  which  extend  over  and  coordinate  them 
so  that  correspondence  is  readily  effected  with  the  changing 
volume  of  the  viscus  ;  otherwise,  the  ureters  should  be  bent 
upon  themselves  during  the  filling,  thereby  interrupting  the 
flow,  while  during  expulsion  they  would  be  subject  to  terrific 
strain,  painful  even  to  contemplation.  Finally,  the  to-and-fro 
movements  in  the  kidneys  themselves  during  respiration, 
from  the  action  in  the  diaphragm,  would  also  call  for  the 
longitudinal  muscles  in  the  ureters,  the  same  as  in  the  oesopha- 
gus, vena  cava,  etc.     Hence  these  muscles. 

Displacement  of  the  kidneys  is  obviated  by  the  layer 
of  peritoneum  that  covers  and  holds  them  against  the  pos- 
terior wall  of  the  abdomen ;  though  occasionally  we  have  "a 


296  TIIE   ACTION   IN    THK   KIDNEYS. 

floating  kidney,"  the  organ  seeming  to  slip  about  too  freely  in 
the  general  cavity,  not  being  securely  held  by  the  overlying 
peritoneum  and  connective  tissue,  forming  a  kind-  of  mesenteric 
ligament  to  the  organ,  which  also  shows  the  necessity  for 
the  special  anatomical  dispositions  in  the  artery,  vein  and 
ureter  for  effecting  elongation  and  contraction  during  respira- 
tion, so  as  not  to  interrupt  the  functions  in  the  organ.  And 
looking  at  all  these  beautiful  adjustments  for  special  work,  can 
it  be  doubted  for  a  single  moment  that  the  arrangements  which 
obtain  in  the  pelvis  of  the  kidney  should  not  expedite  the 
functions  in  these  organs,  whereby  the  whole  is  brought  in 
correspondence  ?  Thus,  in  the  expanded  portion  forming  the 
buccal  projection  of  the  pelvis  (Fig.  118,  6),  the  muscles  are 
thick,  but  thin  in  the  calices,  and  cease  altogether  at  the 
papillae  ;  it  follows  that  expansion  in  the  muscles  must  inevi- 
tably exert  a  suction  force  upon  the  contents  in  the  papillae  and 
uriniferous  tubes.  It  is  all  very  wonderful ;  but  what  more 
wonderful  than  Life  itself  ?  And  when  undifferentiated  proto- 
plasm locomotes  from  place  to  place — moreover,  is  highty  sen- 
sitive— it  looks  still  more  wonderful. 

And  if  the  ureters  should  do  nothing  more  than  elongate  and 
shorten  with  respiration,  and  with  the  filling  and.  the  emptying 
of  the  bladder,  this  of  itself  would  exert  a  suction  force  upon  the 
renal  secretions ;  but  there  is  every  reason  for  believing  in  a 
special  action  in  the  pelvis  of  the  kidney. 

In  order  to  account  for  the  rapid  appearance  of  substances 
in  the  urine,  the  old  authors  conceived  that  a  short  route  ex- 
isted from  the  stomach  to  the  kidneys.  Well,  there  is  a 
short  and  direct  route — as  direct  as  it  can  be  made — but  it  is 
through  the  nervous  channels  that  the  thing  is  effected — 
namely,  by  the  renal  ganglion  (Fig  10y,  G),  and  the  nerves  and 
ganglia  of  the  solar  plexus,  reflected  thence  over  the  coeliac 
axis  (G.  F,  4  ;  G,  C,  4) ;  and  since  sensory  impressions  made 
upon  the  gastric  mucous  membrane  would  tend  to  enlarge  the 
lumen  of  the  renal  arteries  by  reflex  action,  and  thereby 
flush  the  glomeruli  and  increase  the  secretions  in  the  organs 
correspondingly,  we  can  readily  understand  the  rapid  reap- 
pearance of  the  substances  in  the  urine,  especially  when  we 
remember  the  mechanics  in  the  stomach  for  compelling  absorp- 


THE   ACTION   IIST    THE   KIDNEYS.  297 

tion,  and  that  the  entire  circuit  of  the  blood  is  made  within  a 
minute.  In  this  manner,  then,  we  can  readily  understand  the 
specific  action  of  diuretics,  which,  by  acting  upon  the  nerves 
and  ganglion  of  the  renal  plexus,  expands  the  lumen  of  the 
arteries  and  increase  the  blood  supply  to  the  organs  corre- 
spondingly. 

The  rich  nervous  connections  subsisting  between  the  renal 
plexus  and  the  splanchnic  nerves  (Fig.  J  09,  9  ;  G,  A,  b),  the 
third  splanchnic  terminating  in  the  plexus  (7,  S),  will  give 
some  idea  of  the  importance  of  the  renal  functions  for  main- 
taining a  balance  in  the  organism,  since  it  is  through  these 
nerves  that  the  lumen  of  the  artery  is  regulated  ;  also,  why  ap- 
plications made  to  the  skin  surface  in  the  lower  dorsal  and 
upper  lumbar  regions  should  be  reflected  thence  upon  the  renal 
arteries  for  contracting  the  lumen  in  cases  of  engorgement  or 
inflammatory  processes  in  the  organs.  Look  at  it,  please  ;  it 
is  of  great  importance.  The  sensory  impressions  in  the  skin, 
where  the  applications  are  made,  are  transmitted  over  the  rela- 
tive intercostals  to  the  appellate  reflex  centre  in  the  spinal 
cord,  thence  through  the  third  splanchnic  nerve  (7,  8)  to  the 
renal  plexus.  Of  course,  force  is  also  transmitted  through 
the  other  splanchnic  nerves,  but  the  relations  of  the  third 
splanchnic  would  indicate  it  to  be  the  main  channel.  It  is 
needless  to  extend  the  matter. 


CHAPTER  XIII. 

ADJUSTMENTS  IX  THE  WALLS  OF  THE  ABDOMEN  WITH  RESPI- 
RATION  AND   THE   FUNCTIONS    IN   THE   PELVIC   YISCEKA. 

Action  in  the  Mesentery — Ad justments  with  Respiration  and  the  Functions  in  the  Pelvic 
Viscera— Adjustments  -with  Gravitation  and  the  Erect  Position— Formation  of  the 
Sacral  Promontory  in  Man — Adjustments  in  the  External  Oblique  Muscles — Ex- 
ternal Oblique  Muscles  and  Diaphragm  Antagonizing  each  other  in  Respiration — 
The  Respiratory  Plane  upon  which  the  Viscera  Move  in  Respiration — Action  in  the 
Trausversalis — Action  in  the  Internal  Oblique  ;  the  Relations  it  Sustains  to  the 
Pelvic  Viscera — Mode  of  Coordinating  the  Muscles  in  the  Abdomen  with  the  Pelvic 
Viscera — How  the  Diaphragm  Assists  the  Action— Os  Sacrum  and  Sacro-Ischiadic 
Ligaments,  the  Floor  of  Support  to  the  Pelvic  Viscera,  as  also  the  Point  of  Resist- 
ance to  the  Detrusor  Force  in  the  Abdomen — Functions  of  the  Levator  Ani — Me- 
chanics in  Eniesis — The  Stomach  Energetically  Compressed  by  the  Simultaneous 
Contraction  of  the  Diaphragm  and  Muscles  in  the  Abdomen,  Compelling  Rapid  Re- 
gurgitation. 

Frequent  reference  has  been  made  to  the  action  of  the  mass 
of  intestines  known  as  kt  the  mesentery  "  (Fig.  119),  in  connec- 
tion with  respiration  and  circulation  ;  transmitting  the  force 
in  the  walls  of  the  abdomen  upon  the  stomach,  liver,  portal 
and  lower  cava  systems,  spleen,  pancreas,  thoracic  duct,  and 
the  glands  of  the  mesentery  for  increasing  the  action  in  them 
all,  and  compelling  the  venous  blood  and  lymph  within  the 
abdomen  to  flow  into  the  chest-cavity  simultaneously  during 
inspiration,  as  has  already  been  fully  set  forth  ;  and  it  remains 
to  show  the  relations  it  also  sustains  to  the  viscera  in  the  pel- 
vis for  increasing  the  action  in  them,  in  the  bladder  daring 
urination,  and  in  the  colon  and  rectum  during  defecation, 
since  the  action  in  the  walls  of  the  abdomen  is  in  concert 
with  the  action  in  the  pelvic  viscera ;  for  the  parts  are  fully 
coordinated  by  means  of  the  nerves  extending  into  them  from 
the  cerebro-spinal  axis,  and  it  all  performs  as  but  a  single 
organ  only  for  expelling  the  waste  products,  while  the  reflex 
actions  for  producing  it  are  propagated  from  sensory  impres- 
sions in  the  mucous  surface  of  the  special  viscus ;  the  same 


THE     MESEXTERY. 


299 


mechanics  also  applying  for  the  contents  in  the  womb.  This 
mass  of  intestines,  occupying  the  mid -abdominal  regions,  is  in 
a  manner  isolated  by  the  ligament  (Fig.  119,  3),  the  only 
points  of  connection  being  the  duodenum  above,  where  the  je- 
junum (i)  terminates,  and  the  csecal  pouch  below,  where  the 
ileum  ( . )  ends  ;  and  by  simply  dividing  the  intestines  at  these 
two  points,  and  the  ligament  (3)  connecting  it  with  the  lumbar 
vertebrae,  the  mass  is  readily  removed  from  the  abdomen ; 


Eig  119.— The  Mesentery.     1,  1,  1,  Coils  of  jejunum  :  8,  2,  coils  of  ileum  ;  3,  ligament 

of  mesentery. 

while  the  ligament  itself  may  be  spanned  by  the  hand  ;  this, 
notwithstanding  15  to  20  feet  of  intestines  in  the  man,  40  to 
50  in  the  horse,  and  5>U  to  60  in  the  ox,  are  thus  embraced  by 
the  ligament.  Mlrdbile  !  "Well,  by  reason  of  gravitation,  the 
ligament  in  the  quadruped  (Fig.  25,  If)  is  perpendicular,  while 
in  the  erect  position  of  the  trunk,  as  in  man  (Fig.  120),  the  vis- 
cera gravitate  against  the  lower  abdominal  walls  (/,  E)  and  the 
widely-expanded  ilia,  carrying  the  ligament  with  them,  so  that 
it  now  occupies  an  oblique  position  in  the  abdomen  {M,E),  gravi- 
tation compelling  this  circumstance.  Of  course,  the  weight  in  the 
colon  (c),  stomach  (s),  spleen,  and  liver  (L)  is  also  sustained  bv 


BOO 


POSITION"   OF   'I  HE   VISCERA    IN   THE   ABDOMEN. 


the  mesentery,  but  transmitted  of  necessity  to  the  walls  of  the 
abdomen  and  pelvis  as  the  common  floor  of  support,  since  it 
would  not  do  to  have  them  sagging  to  the  diaphragm  or  any 
strain  to  the  ligaments  ;  otherwise  inevitable.  But  one  thing 
calls  for  another.     Thus,  for  obviating  the  pressure  in  the  pelvic 


Fig.  120. — Longitudinal  Section  of  the  Abdomen,  showing  position  of  the  mesentery  with 
the  body  erect,  gravitating  downward  against  the  lower  abdominal  walls  and  false 
pelvis.  M,  center  of  mesentery  ;  J,  /,  cods  of  small  intestine  ;  E,  anterior  wall 
of  the  abdomen  ;  B,  bladder  ;  R,  rectum  ;  C,  colon  ;  S,  stomach  ;  L,  liver  ;  D, 
diaphragm  ;  P,  pancreas  ;  Q,  duodenum. 

viscera  produced  by  the  weight  of  the  viscera,  the  pelvis  is 
tilted  backicard  in  man,  so  as  to  throw  the  mesentery  against 
the  lower  abdominal  walls  (J/,  E)  and  the  expanded  ilia,  a 
few  loose  coils  only  lightly  resting  against  the  pelvic  viscera 
{.R,  B),  whereby  impact  from  the  diaphragm  during  inspiration 
and  from  locomotion,  otherwise  inevitable,  which  would  tend 
to  force  out  the  contents  in  the  organs,  subjecting  them  to 


FORMATION"    OF   THE    SACEAL    PROMONTORY.  301 

strain  as  well,  is  avoided.  It  is  simply  wonderful.  But  there 
is  an  obvious  reason  for  every  one  of  the  adjustments  that 
obtain  in  man.  Please  think  over  it  carefully.  I  can  do  no 
more  to  make  it  plainer.  But  to  me,  however,  it  seems  suf- 
ficiently obvious.  This  backward  tilting  of  the  pelvis,  produc- 
ing the  sacral  promontory  in  man,  is  the  result  of  the  erect 
position  of  the  trunk  and  the  reactionary  forces  in  locomo- 
tion— gravitation,  the  propelling  force  for  driving  the  body 
forward,  together  with  impact  in  the  acetabula  transmitted, 
through  the  crural  bones,  making  them  the  points  of  resist- 
ance, tending  to  carry  the  pelvis  backward  ;  hence,  the  sacral 
promontory  in  man.  It  is  deeply  interesting,  containing  a 
number  of  features  connected  with  the  bones.  It  is  needless  to 
add  that  the  great  development  of  the  pelvic  bones  in  man  has 
similar  explanation,  strain  tending  to  fall  here  during  locomo- 
tion from  the  weight  in  the  body  and  viscera.  *  Before  proceed- 
ing to  the  adjustments  with  the  pelvic  viscera,  however,  it  will 
be  necessary  to  first  consider  the  adjustments  with  respiration, 
since  this  is  fundamental  in  the  organism  ;  when  it  will  be  in 
order  to  regard  the  pelvic  arrangements,  making  it  logical  and. 
scientific,  at  the  same  time  omitting  nothing  which  a  physio- 
logical treatise  should  do,  taking  in  all  the  relative  anatomy 
by  showing  the  adaptations  of  means  to  ends,  since  the  whole 
relates  to  special  work  for  which  they  are  the  relative  adjust- 
ments. 

Action  in  the  External  Obiiqui. — The  external  oblique 
muscles  (one  for  either  side),  obliquus  externus  abdominis,  is 
so  called  from  the  obliquity  of  the  fibres  which  bisect  the 
longitudinal  axis  in  the  body  at  an  acute  angle  (Fig  121).  It 
springs  by  eight  fleshy  digitations  from  the  external  surface 
of  the  eight  inferior  ribs,  and  sweeping  boldly  downward  and 
inward  embraces  the  entire  abdomen  in  front  and  upon  the 
sides,  the  dense  fibrous  leaflet,  in  which  the  muscular  bundles 
terminate,  sweeping  over  the  powerful  rectus  abdominis  (Fig. 
121),  indicated  by  the  band  of  dark  fibres  (linese  transversa) 
to  inosculate  with  the  fibres  from  the  opposite  side  in  the  linea 
alba,  whereby  the  anterior  portions  of   the  abdomen,   where 


For  full  particulars,  see  work  on  "'  Gravitation  and  Development." 


302 


ACTION    IX   TIIK    I.XTEEXAL    OBLIQL'E   MUSCLES. 


strain  tends  to  fall  from  the  weight  in  the  viscera,  is  greatly 
strengthened ;  at  the  same  time,  this  aponeurosis  subserves 
important  purposes  in  conjunction  with  the  aponeuroses  from 
the  other  muscles,  in  maintaining  the  rectus  abdominis  in  posi- 
tion, forming  a  strong  sheath  for  the  powerful  cable-like  mus- 


Fig.  121. — Showing  direction  of  the  fibres  of  the  external  oblique  muscle. — Gray.     The 

letter  A  is  added. 

cle  extending  from  sternum  to  pubes,  that  it  may  not  slip  out 
of  position  in  the  rude  experiences  to  which  the  animal  is  sub- 
ject ;  also,  tending  to  fly  up  from  its  bed  with  every  contrac- 
tion in  the  muscle,  especially  when  energetic,  as  when  the 
body  is  being  flexed.  Below,  the  fibres  are  inserted  into  the 
anterior  portion  01  the  crest  of  the  ilium  and  pubes,  which 
assist  the  recti  abdominis  and  internal  obliqui  in  flexing  the 
trunk  at  the  ]  umbar  vertebrse  in  bending  over  or  stooping.    Now, 


TJ1E     RESPIRATORY    PLANE.  303 

then,  why  should  the  fibres  in  the  external  muscular  layer 
of  the  abdomen  have  this  oblique  direction,  downward  and 
inward,  embracing  the  abdomen,  since  the  matter  has  relation 
to  special  work  which  these  muscles  perform  and  are  the 
relative  adaptations?  It  is  readily  answered.  When  they 
contract,  they  pull  the  viscera  upward  and  backward,  the 
direction  of  the  force  being  upon  a  plane  extending  from 
the  umbilicus  (a)*  to  the  upper  dorsal  curvature  (A),  which 
is  the  respiratory  plane,  all  the  fibres  running  in  this  direction. 
Look  at  it  well,  please  ;  it  will  pay  you,  for  the  gods  have  left 
their  mark.  Well,  when  these  muscles  pull  the  mesentery  with 
the  adjacent  viscera  toward  the  deeper  portions  of  the  chest  dur- 
ing expiration,  the  diaphragm  expands,  in  order  to  admit  of  the 
movement,  which  produces  high  pressure  in  the  lungs,  while  the 
insertions  into  the  lower  ribs  have  the  effect  of  pulling  them 
downward,  whereby  the  action  in  the  lungs  is  greatly  increased, 
causing  the  air  and  blood  to  rush  out  of  the  alveoli  with  corre- 
sponding energy  But  when  the  action  is  reversed,  this  lets  down 
the  viscera  again,  since  they  must  follow  the  floor  of  support, 
while  the  action  in  the  diaphragm,  by  reason  of  the  insertions 
into  the  ends  of  the  seven  lower  ribs  (Fig.  26)  and  lumbar 
vertebrae,  and  the  manner  it  is  ballooned  in  the  chest-exca- 
vation, throws  the  viscera  forward  and  downward  (downward 
and  backward  in  the  horizontal  position),  pressing  them  against 
the  anterior  abdominal  walls,  while  the  ribs  are  thrown  upward, 
flaring  them  open  upon  the  sides,  caused  by  the  muscular  fibres 
in  the  diaphragm  (Fig.  27,  F,  F)  pulling  upon  the  cartilaginous 
ends  of  the  ribs  in  the  long  axis  (pp.  89-91) ;  hence  this  circum- 
stance. This  action  produces  low  pressure  in  the  chest,  the 
lungs  at  the  same  time  expanding  for  confining  it  to  the 
alveoli ;  whereby  the  air  and  blood  are  compelled  into  the 
chambers  simultaneously.  And  by  this  alternating  action  in 
the  diaphragm  and  muscles  in  the  abdomen  we  have  the  pump- 
ing movements  produced  in  respiration,  the  lungs,  of  course, 
acting  in  concert.  In  this  manner,  then,  the  respiratory  plane 
obviates  impact  in  the  pelvic  viscera  during  inspiration  ;  other- 
wise inevitable,  as  must  appear  obvious. 

*  The  terminal  letter  of  "  linea." 


304  THE   RESPIRATORY   PLATSTE. 

The  wide  fibrous  aponeurosis,  spreading  out  like  a  sheet 
over  the  anterior  portions  of  the  abdomen,  distributes  force 
evenly  over  the  viscera,  so  that  undue  pressure  cannot  fall 
upon  any  one  of  the  organs,  each  one  sustaining  its  relative 
amount  of  pressure  according  to  extent  of  surface.  And  the 
great  role  the  muscles  perform  in  respiration  has  forcible  illus- 
tration by  simply  placing  the  open  hand  upon  the  sides  of  the 
abdomen  when  talking,  whistling  or  singing,  the  faintest  vocal 
resonance  being  promptly  reported  by  the  movements  in  these 
muscles,  while  the  action  swells  with  the  volume  of  the  sound. 
There  is  no  mistaking  it.  And  there  is  nothing  more  perfect 
than  the  arrangements  that  obtain  in  this  respect.  The  rounded 
lower  border  of  the  aponeurosis,  known  as  Poupart's  ligament, 
inserted  into  the  spinous  processes  of  the  ilium  and  pubis, 
spans  the  femoral  artery,  vein,  crural  nerve,  flexor  muscles  of 
the  thigh  (psoas  and  iliacus  internus),  like  a  ligamentum 
arcuatum  in  the  diaphragm,  while  the  powerful  deep  fascia  of 
the  thigh  {fascia  lata)  inserted  into  it  hold  it  iirmly,  so  that 
during  contraction  of  the  muscle  strain  cannot  fall  here.  At 
the  same  time  it  gives  protection  to  the  vessels.  Very  pretty — 
all  of  it.  We  now  pass  to  the  transfer  salts,  the  auxiliary 
muscles  for  increasing  the  action  in  the  obliqui,  which  includes 
the  internal  as  well  as  the  external  obliqui,  the  action  in  both 
sets  of  muscles  receiving  important  aid  from  the  pair  of  muscles 
we  shall  now  bring  before  you. 

TJie  Action  in  tlie  Transfer salts  (lumbo-abdomhialis). — The 
transversalis  (Fig.  12:2)  is  the  innermost  of  the  three  muscular 
layers  to  the  abdomen,  the  peritoneum  covering  it,  and  the 
viscera  lying  against  it.  As  the  name  indicates,  the  fibres  run 
transversely  ;  the  muscular  fibres  spring  from  the  internal  sur- 
face of  the  cartilages  of  the  six  inferior  ribs,  lumbar  fascia,  crest 
of  ilium,  and  Poupart's  ligament ;  while  the  fibrous  aponuerosis 
in  which  the  muscle-fibres  terminate  inosculates  with  that  from 
the  opposite  muscle  in  the  linea  alba,  passing  under  the  rec- 
tus abdominis,  forming  the  inner  layer  of  its  sheath,  save  at 
the  lower  portion,  where  the  fibres  pass  in  front  of  it,  blending 
with  the  fibrous  layer  from  the  internal  oblique  muscle  (Fig.  1 22), 
the  rectus  muscle  having  three  layers  of  fibrous  aponeurosis  at 
this  point  for  supporting  the  muscle,  strain  tending  to  fall  in 


THE  ACTION   IN   THE   TKAXSVKKSALES. 


305 


this  locality  (Fig.  120,  E),  from  the  weight  in  the  viscera ; 
hence  this  circumstance.  And  for  giving  the  transverse  mus- 
cles a  firm  point  from  which  to  contract  upon  the  abdominal 
viscera,  the  posterior  aponeurosis  forming  the  lumber  fascia 
{Fig.  122)  is  divided  up  into  three  broad  leaflets,  the  anterior 


Fig.  122.— Showing  direction  of  the  fibres  of  the  transversalis  muscle.— Gray. 

and  middle  being  inserted  into  the  base  and  apex  of  the  trans- 
verse processes  of  the  lumbar  vertebrse,  and  the  posterior  into 
the  powerful  aponeurosis  of  the  latissimus  dorsi,  so  that  force 
with  security  are  insured.  Now,  then,  contraction  in  the 
transversales  must  inevitably  diminish  the  transverse  axis  in 
the  abdomen  ;  and  should  this  occur  simultaneously  with  con- 
traction in  the  external  obliqui  (which  is  the  case),  it  is  easily 
perceived  that  it  would  assist  the  action  in  these  muscles  cor- 


306       THE    ACTION   IN   THE   INTERNAL   OBLIQUE  MUSCLES. 

respondingly.  It  could  not  do  otherwise,  in  the  very  nature 
of  things.  In  short,  the  whole  anterior  walls  of  the  abdomen 
from  sternum  to  pubes  are  pulled  backward  by  contraction  in 
these  muscles,  thereby  increasing  pressure  in  the  abdomen  cor- 
respondingly for  compelling  the  viscera  toward  the  chest  during 
expiration,  while  the  extensive  insertions  into  the  ribs  should 
keep  them  down  for  increasing  pressure  in  the  lungs  ;  in  this 
way  also  aiding  the  external  obliqui.  Finally,  the  two  pairs 
of  muscles  act  together  and  simultaneously,  the  action  alter- 
nating with  the  action  in  the  diaphragm  ;  and  by  aid  of  these 
muscles  the  mesentery  is  moved  to  and  fro  in  a  muscular 
casket,  as  it  were,  the  great  pendulum  in  the  clock-work,  by 
the  force  in  the  medulla  oblongata.  That  will  answer  for  res- 
piration. Now,  then,  in  regard  to  the  action  in  the  pelvic 
viscera  and  the  mode  of  concentrating  force  in  the  pelvis  in 
connection  with  the  special  functions  in  the  viscera.  Here 
comes  in  the  great  role  in  the  internal  oblique  muscles,  to 
which  the  other  two  pairs,  the  transversales  especially,  are 
auxiliary ;  as  also  the  diaphragm,  the  whole  being  available 
for  concentrating  force  in  the  pelvis. 

Concerning  the  Mechanics  for  Expelling  Waste  Prod- 
ucts in  the  Pelvic  Viscera  and  the  Contents  in  the 
Womb. 

The  disposition  of  force  in  the  walls  of  the  abdomen  for 
compelling  out  the  contents  in  the  pelvic  viscera,  which  func- 
tion as  receptacles  for  waste  products,  notably  the  bladder  and 
rectum,  inclusive  of  the  contents  in  the  womb,  is  also  very  per- 
fect. By  reason  of  density  in  the  fsecal  matter  from  absorp- 
tion of  the  aqueous  portions  in  the  colon,  much  force  is  needed 
for  effecting  expulsion,  which  is  also  available  for  increasing 
the  action  in  the  bladder  and  womb  during  urination  and 
parturition,  the  parts  being  fully  coordinated  with  the  viscus, 
by  means  of  the  nerves  running  into  them  from  the  spinal 
axis,  as  before  remarked  Now,  then,  for  producing  this 
force  we  have  the  following  arrangement  in  the  muscles, 
notably  :  1.  The  powerful  internal  oblique  muscles,  or  obliqui 
internus  abdominis  ascendens  (Fig.  126),  springing  from  the 
middle  of  the  crest  of  the  ilium  for  two- thirds  its  length,  outer 
half  of  Poupart's  ligament  and  lumbar  fascia,  whence  the  fibres 


THE  ACTION   IN   THE   INTERNAL   OBIIQUE  MUSCLES. 


307 


proceed  upward  and  inward,  to  be  inserted  into  the  lower 
border  of  the  four  inferior  ribs,  and  the  whole  of  the  linea 
alba  from  the  ensiform  cartilage  to  the  pnbes  by  means  of  the 
broad  aponeurosis,  where  the  fibres  inosculate  with  those 
from  the  opposite  side      Along  the  border  of  the  upper  three- 


Vortjoincit  Tent/on 
CREMA5TER- 


Fig.  123.— Showing  direction  of  the  fibres  of  the  internal  oblique  muscle.— Gray. 

fourths  of  the  rectus  abdominis  the  aponeurosis  splits  into 
two  layers  for  embracing  the  muscle,  forming  the  middle  layer 
of  its  sheath,  but  at  the  lower  portion  it  again  comes  in  front 
of  the  muscle,  following  the  course  of  the  transversalis  (Fig. 
122),  and  for  the  same  reason;  near  Poupart's  ligament  the 
fibres  come  downward  over  the  spermatic  cord  to  be  inserted 


308       THE    ACTION    IN    THE   INTEFrXAL    OBLIQUE    MUSCLES. 

into  the  pubes.  That  will  serve  the  purpose  of  description. 
Now,  then,  what  can  be  the  purpose  of  this  arrangement 
in  the  fibres  of  the  internal  oblique  muscle,  if  it  be  not 
for  increasing  pressure  in  the  pelvis  in  connection  with  the 
special  functions  in  these  organs  %  At  the  same  time,  they 
also  assist  in  flexing  the  trunk  at  the  lumbar  vertebrae, 
as  well  as  the  expiratory  effort  by  pulling  the  ribs  down- 
ward during  this  time,  and  when  the  action  is  energetic 
they  pull  vigorously  npon  them.  But  the  major  number  of 
the  fibres  extend  out  over  the  walls  of  the  abdomen,  and  by 
means  of  the  dense  aponeuroses  the  whole  mesentery  is  em- 
braced, and  when  contracting  vigorously,  as  in  passing  the 
faecal  contents,  this  is  pulled  into  the  excavation,  pressed, 
piston-like,  down  upon  the  pelvic  viscera  for  compelling  out 
the  contents  in  the  rectum  or  bladder,  as  the  case  may  be. 
First,  the  body  is  flexed  or  bent  upon  itself  (Fig.  124),  for 
shortening  the  longitudinal  axis  in  the  abdomen,  which  is 
accomplished  by  the  action  of  these  muscles  and  the  recti 
abdominis,  at  the  same  time  the  thighs  are  flexed  by  the 
action  of  the  psora  and  iliaci  interni,  which  brings  the  long 
axis  in  the  pelvis  in  correspondence  with  the  long  axis  in  the 
abdomen,  whereby  the  force  in  the  diaphragm  is  brought 
to  bear  upon  the  pelvic  viscera.  Everything  being  ready, 
the  diaphragm  contracts  by  inspiratory  effort  for  increasing 
pressure  in  the  abdomen  ;  finally,  the  internal  obliqui  are  put 
into  action,  which  force  the  mesentery  duwnioard  into  the 
excavation  (Fie;.  124,  31),  the  loose  coils  of  intestine  glid- 
ing readily  over  one  another  under  the  force  in  the  abdomen, 
downward  and  backward  against  the  rectum  (i?),  which  can- 
not escape  and  must  endure  the  pressure,  at  the  same  time 
contracting  energetically  for  assisting  the  action,  while  the 
sphincters  expand  simultaneously  for  reducing  resistance,  as 
also  for  obviating  strain  and  rude  friction  in  the  parts,  otherwise 
inevitable.  It  is  simply  perfect.  As  will  be  seen,  the  ligament 
is  carried  downward,  and  is  no  longer  upon  the  plane  it  occu- 
pies in  the  erect  position  or  toward  the  anterior  abdominal 
walls  (Fig.  120,  31,  E).  And  by  thus  pulling  the  mesenteric 
piston  with  great  force  down  upon  the  bladder  and  rectum, 
together  with  the  action  in  the  viscus,  the  contents  are  ex- 


CONCENTRATING  FORCE   IN   THE   PELVIC   VISCERA.         309 

pelled.  In  short,  nothing  could  be  more  admirable  than  the 
arrangements  which  obtain  for  increasing  pressure  in  the 
pelvic  excavation  commensurate  with  the  functions  in  these 
organs,  the  same  remark  applying  for  every  stage  in  develop- 
ment.   Thus  in  the  quadruped  the  lumbar  vertebrae  are  flexed 


Fig.  134.— Longitudinal  Section  of  the  Abdomen,  showing  position  of  the  mesentery 
during  expulsion  of  waste  products.  M,  middle  of  mesentery  ;  I,  I,  coils  of  small 
intestine  ;  E,  anterior  abdominal  wall  ;  B,  bladder  ;  R,  rectum  ;  C,  colon  ;  S, 
stomach  ;  L,  liver  ;  D,  diaphragm  ;  Q,  duodenum  ;  P,  pancreas. 

by  incurvating  them  upward,  which  is  done  by  contracting 
the  recti  and  obliqui,  as  in  man,  the  muscles  upon  the  dorsal 
surface  at  the  same  time  expanding  or  elongating  pari  passu 
with  this  movement.  Then  the  diaphragm  is  contracted  for 
shortening  the  longitudinal  axis  ;  finally,  the  transversales 
and  internal  oblique  muscles  are  energetically  contracted  for 


:*10         CONCENTRATING   FOllCE   IN   THE  PELVIC   VISOKRA. 

shortening  the  transverse  axis  and  compelling  the  mesenteric 
pi  stun  into  the  pelvic  excavation. 

The  action  is  studied  to  great  advantage  in  the  dog  and  cat, 
in  which  the  excrement  is  hard,  and  Unusual  force  is  needed 
for  expelling  it.  It  goes  hard  with  them,  and  they  work  away 
vigorously  in  performing  it,  concentrating  their  force  in  the 
pelvic  viscera,  the  rectum  especially,  greatly  arching  the  spine 
upward  and  bringing  the  force  in  the  diaphragm  and  abdominal 
muscles  to  bear  upon  the  rectum  transmitted  through  the 
mesenteric  piston.  This  condition  of  the  rectal  contents  would 
explain  the  great  development  of  the  muscles  in  this  locality, 
which  are  much  more  numerous  than  in  any  other  portion  of 
the  canal. 

In  the  bladder,  on  the  contrary,  the  contents  are  liquid ; 
hence,  not  so  much  force  is  needed  for  expelling  them,  unless, 
forsooth,  there  is  obstruction  to  the  natural  now — e.  g.,  stric- 
ture— when  the  walls  are  hypertrophied,  becoming  extremely 
thick  in  course  of  time.  When  the  animal  desires  to  pass 
urine,  he  puts  himself  into  a  proper  position,  when  the  me- 
chanics is  set  in  motion  for  effecting  expulsion,  all  the  parts 
being  coordinated  by  the  central  nervous  system,  so  that  the 
whole  performs  as  but  a  single  organ  only,  as  in  the  case  of  the 
rectum. 

In  the  parturient  woman,  this  reflex  action,  which  is  propa- 
gated from  the  viscus  through  the  spinal  cord,  is  forcibly 
illustrated  by  simply  drawing  upon  the  perineum  by  means 
of  the  index  and  middle  fingers  inserted  between  the  labia, 
when  a  tremendous  expulsive  effort  at  once  sets  in,  the  patient 
contracting  the  diaphragm  and  holding  her  breath,  then  power- 
fully contracting  the  internal  oblique  and  transversales,  the 
womb  at  the  same  time  contracting,  all  the  parts  acting  in 
concert,  as  in  the  other  cases,  the  cervix  at  the  same  time  ex- 
panding. 

In  the  case  of  birds  the  legs  are  brought  further  under  the 
body,  which  is  tilted  a  little  up  anteriorly  and  lowered  pos- 
teriorly (which  favors  gravitation) ;  at  the  same  time,  the  abdo- 
men and  cloaca  are  contracted  for  expelling  the  contents,  faecal 
matter  or  ovum,  as  the  case  may  be  the  principle  being  the  same. 
For  expelling  the  ovum  great  force  is  needed,  while  time  is 


OS   SACEUM   THE   TEUE  FLOOR   OF  THE    PELVIS.  311 

required  for  effecting  the  requisite  expansion  in  the  sphincters. 
The  event  is  announced  with  great  joy.  In  all  the  mechanics  it 
is  fundamentally  the  same,  requiring  the  action  in  the  mesen- 
tery and  the  muscles  in  the  abdomen,  in  connection  with  the 
action  in  the  special  viscus  for  effecting  expulsion  of  the  con- 
tents. 

Finally,  we  have  to  note  several  adjustments  for  conserving 
structure  and  increasing  function,  charming  to  look  upon  ;  nota- 
bly :  a.  The  incurvation  of  the  sacrum,  which  is  bent  in  both 
the  longitudinal  and  transverse  axes,  or  dished,  so  to  speak,  in 
two  directions,  the  rectum  occupying  the  long  diameter,  and  by 
means  of  the  articulations  with  the  ilia  is  inclined  at  an  ob- 
lique angle  with  the  spine ;  in  other  words,  tilted  backward 
in  the  upper  portions ;  but  being  bent  upon  itself,  the  lower 
portion  is  again  brought  forward,  across  the  longitudinal 
axis  in  the  pelvis,  and  forming  with  the  sacro-ischiadic  liga- 
ments (Fig.  125,  10,  9)  the  true  floor  of  the  pelvis  for  sus- 
taining the  weight  in  the  viscera  and  the  force  in  expulsion, 
the  mesenteric  piston  being  driven  plump  up  against  it  with 
the  rectum  sandwiched  between  them  (Fig.  1^4,  M,  i,  R), 
thereby  saving  the  perineum  The  rectum,  thus  lying  in  the 
hollow  of  the  sacrum  through  its  whole  length,  sweeps  out  over 
the  end  of  the  coccyx  to  reach  the  external  surface  to  form  the 
anal  opening.  It  is  easily  retained  in  position  by  means  of 
the  overlying  peritoneum  and  connective-tissue  fibres,  while 
the  terminal  end  is  supported  by  the  broad  muscular  leaflets 
of  the  levator  ani  (Fig.  1 25,  7),  proceeding  from  the  sides  of 
the  rectum,  to  be  extensively  inserted  into  the  adjacent  sides 
of  the  pelvis,  forming  the  portion  of  the  floor  corresponding 
with  the  perineum  ;  and  supporting  the  bladder  and  vagina  as 
well.  It  results,  from  this  arrangement  in  the  parts,  that  but 
little  of  the  detrusor  force  in  the  abdomen  falls  directly  upon 
the  perineum,  the  sacrum  sustaining  it ;  at  the  same  time,  it 
is  made  more  effective  upon  the  faecal  matter,  the  rectum,  of 
course,  contracting  simultaneously  for  increasing  the  action, 
while  the  levator  ani  are  in  concert  so  as  to  obviate  strain  to 
the  perineum,  the  parts  being  fully  coordinated. 

The  manner  Nature  has  solved  this  difficult  problem  in 
mechanics,  which  is  truly  wonderful,  must  be  set  down  to 


812  OFFICE    (»F    THE     LEVATOKES   ANT. 

necessity,  and  as  a  prerequisite  to  the  erect  position,  gradually 
brought  about  in  connection  with  the  other  changes  in  the 
pelvic  framework. 

The  following  illustration  (Fig.  126)  will  show  the  arrange- 
ment and  distribution  of  the  spinal  nerves  to  the  chest  and 
walls  of  the  abdomen.  As  will  be  seen,  the  nerves  to  the 
abdominal  walls  are  nearly  all  intercostals,  so  called,  save  the 
ilio-hypogaster  (1G)  or  first  lumbar  nerve,  to  the  inguinal 
regions,  but  in  the  abdomen  the  intercostals  are  greatly 
enlarged  in  correspondence  with  the  amount  of  muscles  for 
evolving  the  force  in  the  walls  in  connection  with  respira- 


Fig.  125. — Muscles  of  the  Perineum. — Wilson  and  Buchanan. 

tion  and  the  functions  in  the  viscera.  And  one  may  readily 
perceive  how  the  action  in  the  abdomen  may  be  made  to 
alternate  with  the  action  in  the  diaphragm  in  respiration, 
since  the  phrenic  nerves  are  correlated  with  them  in  the 
respiratory  centre ;  as  also  how  the  viscera  are  brought 
into  correspondence  with  the  walls  through  the  reflex  actions 
set  up  by  means  of  the  pneumogastric  and  splanchnic  nerves, 
correlated  with  the  intercostal  and  phrenic  nerves.  Finally, 
how  sensory  impressions  in  the  pelvic  viscera  should  con- 
centrate the  force  in  the  pelvic  excavation,  in  connection 
with  the  special  functions  in  these  organs,  while  the  enormous 
web  of  nerves  and  nervous  ganglia  in  the  hypogastric  plexus 
(Fig.  116),  together  with  the  nerves  running  into  it  and  the 
special  viscera  from  the  spinal  axis,  notably  from  the  fifth 
lumbar  and  sacral  nerves  (Ibid),  would  make  us  readily  under- 


NERVES   OF    THE    ABDOMINAL   WALL. 


313 


stand  how  the  vise  as  itself  should  be  influenced  by  the  force 
from  the  central  nervous  system,  at  the  same  time  it  effects 
the  actions  in  the  abdominal  walls,  all  the  parts  acting  in 
concert  to  this  end.     It  is  complex,  but  readily  understood. 


Fig.  126.— The  Nerves  of  the  Abdominal  Wall  (from  Hirsehfeld  and  Leveille").  1,  Pec- 
toralis  major  (cut)  ;  2,  serratus  magnus  ;  3,  latissimus  dorsi ;  4,  intercostal  muscles  ; 
5,  rectus  abdominis  ;  6,  section  of  obliquus  externus  ;  7,  obliquus  internus  ;  8,  trans- 
versalis  abdominis  ;  9,  9,  ninth  dorsal  nerve  ;  10,  10,  tenth  dorsal  nerve  ;  11,  11, 
eleventh  dorsal  nerve  ;  12,  12,  twelfth  dorsal  nerve  ;  13,  lateral  cutaneous  branch 
of  first  lumbar  (iho -hypogastric)  ;  14,  anterior  cutaneous  branch  of  i  Ho -hypogastric  ; 
15,  anterior  cutaneous  branch  of  ilio-inguinal  ;  16,  ilio-hypogastric  and  ilio-inguinal 
nerves  ;  17,  lateral  cutaneous  branch  of  second  intercostal  nerve  ;  18,  lateral  cuta- 
neous branch  of  intercostal  nerve. 

Concerning  the  Action  in  Enusis  — The  action  in  emesis  is 
also  easily  understood.  Thus,  in  place  of  alternating  in  their 
action,  as  in  the  usual  way  in  respiration,  the  muscles  in  the 


314  THE   ACTION   IN    EMESIS. 

abdomen  and  diaphragm  contract  simultaneously  and  the 
stomach,  caught  between  these  two  forces,  as  in  the  grip  of  a 
vis^\  is  compelled  to  yield  up  its  contents,  which  is  by  re- 
gurgitation ;  for  the  enormous  pressure  within  the  abdomen 
precludes  it  in  the  other  direction,  flowing  from  high  to  low 
pressure,  in  conformity  with  organic  law.  This  would  ex- 
Fig.  127.  Fig  128. 


Fig.  127. — Longitudinal  Section  of  the  Abdomen,  showing  position  of  the  mesentery. 
Fig.  128. — Longitudinal  Section  of  the  Abdomen,  showing  position  of  the  mesentery 

during  vomiting. 

plain  why  emesis  may  be  produced  when  a  bladder  containing 
water  is  substituted  for  the  stomach  by  irritation  propagated 
from  the  pharynx,  which  is  obvious  enough.  But  here,  as 
elsewhere,  the  viscus  acts  in  concert  with  the  muscles  in  the 
abdomen,  contracting  vigorously  upon  the  contents,  as  in  the 
case  of  the  rectum,  bladder  and  womb,  producing  a  reverse 
peristalsis,  since  this  would  assist  the  action  ;  for  Nature  con- 
centrates all  her  available  force  in  expulsive  efforts  The 
short,  loud  inspiratory  sound  characteristic  of  vomiting  is 
produced  by  energetic  inspiration  suddenly  arrested  by  the 
stronger  action  in  the  abdominal  muscles.  The  above  illus- 
trations will  serve  for  impressing  the  matter  (Figs.  127,  128). 


TBE   ACTION  IN    EMESIS. 


315 


Thus  the  mesentery  is  forcibly  compressed  against  the 
stomach  under  the  energetic  action  of  the  muscles  in  the 
abdomen,  the  contracted  diaphragm  being  the  point  of  re- 
sistance, since  both  act  in  concert ;  while  the  correlation  of 
the  pneamogastric,  phrenic,  and  intercostal  nerves  in  the  me- 
dulla oblongata  enables  this  concert  of  action  in  the  stomach, 
diaphragm,  and  the  muscles  in  the  abdomen  to  be  produced. 
The  irritation  is  propagated  from  sensory  impressions  in  the 
gastric  mucous  membrane,  which  sets  up  the  action  in  the  me- 


Fig.  129. — Transverse  Section  of  the  Muscles  in  the  Abdominal  Walls. — Gray. 

dulla  whence  it  is  reflected  over  the  structures.  It  is  all  very 
simple  and  easily  understood.  The  extreme  pressure  upon  the 
liver  and  gall-bladder  would  also  account  for  the  escape  of  the 
bile. 

The  above  instructive  diagram  (Fig.  129)  will  show  the 
relative  position  of  the  muscles  in  the  abdominal  walls  and 
the  manner  the  recti  abdominis  are  kept  in  position  by  the 
strong  fibrous  aponeuroses.  In  the  quadruped  the  weight  of 
the  viscera  is  largely  sustained  by  these  powerful  muscular 
"beams ;  at  the  same  time,  their  aid  is  readily  invoked  for  assist- 
ing in  flexing  the  spine  at  the  lumbar  vertebrae,  for  which  there 
is  frequent  occasion  in  connection  with  the  functions  in  the 
pelvic  viscera,  as  before  remarked.  In  man,  still  more  fre- 
quently for  effecting  the  bending  and  stooping  posture. 


CHAPTER    XIV. 

CIRCULATION    IN    THE    EMBRYO,     MECHANICAL    PRINCIPLE    IN. 

The  Embryo,  an  Aquatic  Animal,  being  Submerged  in  the  Liquor  Amnii,  Deeply  Buried 
in  the  Maternal  Tissues,  and  Sustaining  Itself  the  same  as  the  Fishes,  only  that  a 
Special  Arrangement  Obtains  Respecting  it,  by  Means  of  which  it  both  Respires  and 
Feeds  at  One  and  the  Same  Time  in  the  Maternal  Blood — Placental  Souffle  the 
Analogue  of  Respiration,  the  Rhythms  being  as  One  to  Four  of  the  Foetal  Heart,  or 
the  Same  as  Obtains  in  the  Lungs  and  Heart  in  the  Air-Breather — Mode  of  Connect- 
ing the  Maternal  aud  Foetal  Circulations — The  Pumping  Action  in  the  Placenta 
Aspirating  the  Nutritive  and  Force-Producing  Elements  in  the  Blood  in  the  Sinuses 
and  Expelling  "Waste  Products — Rhythmical  Expansions  and  Contractions  Taking 
Place  in  the  Womb  and  Placenta,  but  which  Alternate  with  Each  Other,  so  that 
when  the  Placenta  is  Expanding,  the  "Womb  is  Contracting,  and  vice  versa,' 
whereby  the  Blood  in  the  Sinuses  is  Rapidly  Renewed,  and  Absorption  Increased 
Correspondingly — Nerves  for  Controlling  the  Arterial  Feeders  to  the  "Womb  for 
Making  Circulation  in  the  Sinuses  Commensurate  with  the  Physiological  Require- 
ments in  the  Embryo,  and  for  Producing  Correspondence  throughout,  so  that  the 
Pumping  Action  in  the  Uterus  and  its  Contents  is  Made  Universal,  as  in  the  Air- 
Breather— Rhythmic  Centre  for  the  Pumping  Actions  Located  in  the  Lumbar  En- 
largement of  the  Spinal  Medulla — The  Rhythmic  Expansions  and  Contractions  in 
the  "Womb  Contrasted  with  the  Action  in  the  Amnion  of  the  "  Chick,"  which  is 
Rocked  to  and  fro  in  the  Egg-Shell  by  the  Rhythmical  Expansions  and  Contrac- 
tions in  the  Amnion  for  Effecting  the  Requisite  Changes  of  Pressure  in  the  Embryo 
and  Allantois  for  Increasing  Circulation,  Making  it  Commensurate  with  the 
Physiological  Requirements. 

This  brings  us  to  circulation  in  the  embryo.  We  have  seen 
that  the  animal  organism  is  based  upon  pressure  and  the  power 
of  producing  rapid  rhythmical  changes  in  pressure  for  increas- 
ing circulation  commensurate  with  the  physiological  require- 
ments, otherwise  impossible  ;  that  respiration  and  circulation 
necessarily  form  a  connected  movement  for  producing  an 
uninterrupted  and  continuous  current  of  the  fluids  between 
the  cell-brood  and  environment  from  which  the  supplies  are 
obtained,  and  into  which  the  waste  products  are  returned  for 
redistribution.  We  now  follow  this  matter  and  take  up  circu- 
lation in  the  embryo,  in  order  to  give  the  true  interpretation 
of  the  phenomena,  anatomical  and  physiological,  appertaining 


THE   EMBRYO   AN  AQUATIC   ANIMAL. 


317 


to  it  also.  There  is  increasing  complexity,  but  from  the  stand- 
point afforded  by  this  law  for  the  circulation  it  is  readily 
explained,  so  that  a  mental  picture  may  be  formed  by  aid  of 
illustrative  diagrams  and  the  special  anatomy.  But  we  must 
begin  with  the  special  environment  and  the  mode  of  maintain- 
ing existence.  After  all,  the  mammilian  is  amphibious,  the 
earlier  portions  of  its  life  being  passed  under  water,  since  the 
embryo  (Fig.  130)  is  submerged  in  the  liquor  amnii  and  deeply 
buried  in  the  maternal  tissues,  while  the  arrangements  that 


Fig.  130. — Gravid  Human  Uterus  and  Contents,  showing  the  relations  of  the  cord, 
placenta,  membranes,  etc.,  about  the  end  of  the  seventh  month. — Dalton.  1,  Decidua 
vera  ;  2,  decidua  refiexa  ;  3,  chorion  ;  4,  amnion. 

obtain  for  producing  growth  and  evolving  force  are  funda- 
mentally the  same  as  in  the  fishes,  due  allowance  being  made 
for  the  special  environment  ;  the  one  feeding  and  resj)iring  in 
the  sea,  the  other  in  the  maternal  blood  by  means  of  a  com- 
posite organ  known  as  the  placenta,  feeding  and  respiring  at 
the  same  time  through  the  placental  organ,  which  thus  sub- 
serves a  double  function,  like  the  intestine  in  the  early  stages 
in  development,  the  animals  feeding  and  respiring  through  it. 
And,  indeed,  a  remnant  of  it  is  still  seen  in  mammalia  in  the 
consumption  of  air  and  absorption  of  oxygen  in  the  stomach 
when  feeding. 

The  physical  conditions  under  which  the  embryo  is  evolved 


318  THE    EMBRYO   AN   -AQUATJC    ANIMAL. 

determine  the  special  vascular  arrangements  for  effecting  cir- 
culation, while  the  transformations  which  accompany  this 
provide  for  the  radical  physical  changes  which  are  ushered  in 
at  the  end  of  the  term,  when  it  becomes  an  air-breather,  the 
fundamental  circumstance  underlying  it  all  being  art  adjust- 
ment with  pressure,  and  the  power  of  effecting  rapid  rhyth- 
mical changes  in  pressure,  for  compelling  circulation  in  the 
measure  of  the  physiological  requirements.  For  example,  we 
have  seen  that  the  rhythmical  expansions  and  contractions  per- 
vading the  body  in  the  air-breather,  and  known  as  respiration, 
compel  oxygen  and  aliment  into  the  circulatory  apparatus  for 
evolving  force  and  producing  growth ;  so,  in  like  manner,  a 
similar  necessity  exists  in  the  embryo  for  compelling  the 
nutritive  and  force-producing  elements  into  its  circulation  for 
producing  growth  and  evolving  force,  which  is  principally 
expended  in  elaborating  its  structures.  But  since  the  embryo 
feeds  in  the  uterine  sinuses  from  which  the  commerce  is 
obtained,  and  into  which  the  waste  products  are  returned, 
this  calls  for  the  differentiation  of  the  placenta  organ,  which 
answers  to  the  more  highly  differentiated  lungs  and  in- 
testinal canal  which  are  to  substitute  it  at  the  end  of 
the  intra-uterine  term  as  the  relative  adjustments  with  the 
larger  environment,  lower  pressure,  and  higher  order  and 
.amount  of  work  which  this  involves.  The  placental  souffle, 
then,  which  is  distinctly  heard  through  the  maternal  struc- 
tures, is  the  analogue  of  respiration  in  the  air-breather,  the 
relative  ratio  of  the  movements  to  the  pulsations  in  the  foetal 
heart  being  also  the  same,  or  as  1  to  4  of  the  latter,  while 
the  villi  are  the  analogues  of  the  villi  in  the  intestinal  canal, 
the  one  submerged  in  the  sinuses,  the  other  in  the  juices  in 
the  intestines.  Since  the  pumping  action  in  respiration  is 
absolutely  essential  for  compelling  the  commerce  into  the  ves- 
sels, it  follows  that  this  circumstance  should  be  represented  in 
the  embryo,  as  the  maternal  blood  does  not  enter  the  embryo, 
the  latter  feeding  out  of  the  sinuses  simply  by  means  of  the 
villi  in  the  placenta  and  this  pumping  action  spoken  of  ;  for 
here,  as  elsewhere  in  the  body,  there  are  no  means  for  increas- 
ing circulation  but  by  rhythmical  changes  in  pressure.  The 
result  must,  then,  be  the  action  in  the  placenta  simulating  res- 


PLACENTAL  SOI  FFLE  THE  ANALOGUE  OF  EESPIEATIOJST.    319 

piration.  The  relative  frequency  of  this  movement  to  the 
action  taking  place  in  the  foetal  heart  is  as  1  to  4,  or  the  same 
as  in  respiration,  as  before  remarked. 

Thus,  in  the  case  of  the  placental  souffle  it  is  30  to  35,  and  in 
the  foetal  heart  the  pulsations  are  from  120  to  140  per  minute  ; 
while,  in  the  case  of  the  air-breather,  the  pumping  action  in 
the  trunk  or  respiration  is  from  16  to  20,  and  in  the  heart  from 
GO  to  J-0  per  minute.  Again,  this  action  in  the  placenta  serves 
not  only  to  pump  the  fluids  into  and  out  of  the  sinuses,  but  at 
the  same  time  it  also  aspirates  the  venous  blood  in  the  embryo 
for  effecting  oxygenation  in  it  the  same  as  obtains  in  the 
lungs  ;  the  heart  and  vessels  assisting  in  the  one  as  well  as  in 
the  other,  since  it  all  forms  a  connected  movement.  We  now 
see  that  by  reason  of  the  great  increase  in  pressure  that  obtains 
in  the  embryo,  the  action  in  the  organs  for  changing  pressure 
is  materially  assisted,  since  the  fluids  flow  more  readily  in  con- 
sequence. And  here  comes  in  the  benefit  of  the  amniotic  fluid, 
which  not  only  increases  pressure  in  proportion,  but  at  the 
same  time  it  serves  to  transmit  the  force  in  the  placenta  and 
uterine  walls  upon  the  embryo  for  compelling  the  blood  to  and 
from  the  sinuses  for  respiratory  purposes,  by  first  increasing, 
then  diminishing  pressure  in  the  embryo,  which  the  actions 
in  the  womb  and  placenta  must  inevitably  effect. 

Thus,  when  the  placenta  expands  for  aspirating  the  fluids  in 
the  uterine  sinuses,  the  organ  advances  into  the  uterine 
cavity,  it  swells  out  and  occupies  more  room,  in  consequence ; 
and,  by  thus  encroaching  upon  the  embryonic  area,  it  produces 
corresponding  increase  in  pressure  upon  the  liquor  amnii  and 
embryo,  with  low  pressure  in  itself,  which  fulfills  the  con- 
ditions for  increasing  circulation  from  the  embryo  to  the 
placenta,  at  the  same  time  that  it  should  aspirate  the  fluids  in 
the  uterine  sinuses.  It  could  not  do  otherwise,  in  the  very 
nature  of  things.  But  during  contraction  in  the  placenta,  the 
opposite  conditions  should  obtain,  since  this  would  determine 
high  pressure  in  the  latter  with  low  pressure  in  the  embryo, 
the  blood  in  consequence  flowing  through  the  umbilical  vein 
with  augmented  speed,  for  the  reason  that  contraction  should 
reduce  the  volume  of  the  placenta,  which  would  inevitably  re- 
duce pressure  in  the  embryo  in  proportion,  the  blood  flowing 


320   PLACENTAL  SCUFFLE  THE  ANALOGTE  OF  EESPIRATION. 

from  one  into  the  other  in  conformity  with  organic  law.  To 
this,  again,  must  be  added  the  action  in  the  foetal  heart  for 
aspirating  the  blood  in  the  placenta.  The  amniotic  media  in 
which  the  animal  lives  obviates  the  necessity  for  the  extensive 
arrangements  for  reducing  pressure  in  the  chest,  which  obtain 
in  the  lighter  media  of  the  atmosphere,  the  heart,  together 
with  the  force  in  the  placenta  and  umbilical  vein,  being  suffi- 
cient for  the  purpose. 

A  subaquatic  existence  calls  for  but  slight  reduction  in 
pressure  in  order  to  compel  circulation  ;  accordingly,  we  have 
the  blood  rushing  into  and  out  of  the  foetal  heart  as  a  result  of 
the  rhythmical  expansions  and  contractions  taking  place  in 
this  organ,  the  same  as  in  the  fishes,  the  principle  being  the 
same  in  this  organ  ;  but  for  increasing  the  action,  the  heart  and 
vessels  are  more  muscular.  The  right  ventricle  is  thicker  and 
stronger  thanilie  left,  as  the  relative  physiological  adjustment 
with  this  mechanics,  since  the  force  in  the  diastole  is  increased 
correspondingly  for  aspirating  the  blood  in  the  umbilical 
vein  and  venous  system,  while  the  force  in  the  systoles  should 
have  similar  increase  for  compelling  the  blood  through  the 
umbilical  arteries  to  the  placental  tufts  and  sinuses,  the  womb 
and  placenta  at  the  same  time  assisting  these  actions  in  the 
manner  as  stated,  the  whole  forming  a  connected  movement 
the  same  as  in  respiration,  since  there  can  be  no  doubt  that  the 
principle  is  the  same  in  both,  the  vascular  apparatus  and  the 
heart  being  coordinated  with  the  rjlacenta  and  womb,  the  same 
as  in  the  lungs  and  muscular  envelope  in  the  air-breather,  else 
the  relative  ratio  of  the  movements  would  be  meaningless. 

By  means  of  this  combined  action  in  the  womb  and  placenta, 
heart  and  vessels,  together  with  the  high  pressure  that  obtains 
in  the  womb,  a  rapid  circulation  is  readily  effected  in  the 
embryo  and  sinuses ;  but  anything  which  should  reduce  pres- 
sure— e.  g  ,  escape  of  the  amniotic  fluid— would  promptly 
destroy  life.  The  lividity  of  the  skin,  which  occurs  in  these 
cases,  proves  conclusively  the  existence  of  venous  stasis  in  the 
systemic  capillaries,  and  insufficiency  of  the  heart's  action  to 
carry  on  circulation  in  the  absence  of  the  normal  pressure 
upon  the  embryo.  How  otherwise  account  for  this  circum- 
stance, since  the  vascular  connections  are  uninjured?    More- 


PLACENTAL  SOUFFLE  THE  ANALOG  *  E  OF  RESPIRATION.    321 

over,  trie  same  circumstance  occurs  to  the  air-breather  when 
pressure  is  too  greatly  reduced,  as  when  suddenly  carried  to 
too  great  an  altitude  in  the  balloon,  the  skin  becoming  livid 
from  venous  stasis  in  the  systemic  capillaries,  producing  in- 
sensibility and  death,  as  occurred  in  the  celebrated  case  at 
Paris,  in  wdiich  the  voyage  was  made  by  two  persons,  the  one 
losing  his  life,  the  other  unconscious  when  the  balloon 
descended  ;  this  notwithstanding  the  extensive  arrangements 
for  changing  pressure  in  the  chest,  which  exist  in  the  latter ; 
and  in  persons  ascending  mountain  ranges,  respiration  and 
circulation  become  more  and  more  embarrassed  as  the  journey 
is  proceeded  with,  and  •  venous  stasis  more  and  more  con- 
spicuous, till  the  limit  of  endurance  is  reached  or  life  itself  is 
terminated.  In  short,  animal  life  has  adjustment  with  press- 
ure, and  whether  in  the  air-breather  or  in  the  embryo,  itself, 
the  balance  must  not  be  too  greatly  disturbed,  else  life  would 
have  a  speedy  termination. 

Such,  in  brief,  is  the  principle  in  the  embryonic  circulation ; 
but  in  order  to  make  the  matter  fully  intelligible  and  easily 
understood,  it  will  now  be  necessary  to  go  briefly  over  the  for- 
mative changes  till  the  animal  is  compelled  into  the  great 
environment  and  becomes  an  air-breather,  the  same  as  the  pa- 
rent. Commencing  with  the  matrix  or  mucous  membrane  of  the 
womb,  we  have,  then,  a  dense  mass  of  uterine  follicles  or  tubu- 
lar glands  (Fig.  131),  packed  closely  together,  the  orifices 
opening  upon  the  mucous  surface  (</),  the  blind,  convoluted 
ends  against  the  muscular  walls  of  the  womb,  and  composed  of 
columnar  epithelium  (Fig.  132).  When  the  impregnated  ovum 
enters  the  uterine  cavity  from  the  Fallopian  tube,  it  pushes  the 
deciduous  membrane  before  it,  and  comes  into  immediate  con- 
tact with  the  congested  and  swollen  mucous  membrane  produced 
by  the  physiological  changes  brought  about  by  impregnation, 
all  the  parts  being  in  correspondence  from  the  action  of  the 
special  nervous  forces  in  the  organs,  the  decidual  membrane 
forming,  as  a  result  of  the  nutritive  changes  set  up  in  the  mu- 
cous membrane  for  feeding  the  ovum  during  the  early  period 
before  intimate  attachment  is  formed  with  the  womb,  and 
enabling  it  to  develop  the  tufts  in  the  chorion  for  that  pur- 
pose.    Now,  then,  when  the  ovum  (Fig.   "J  33)  comes  into  the 


322  PHYSIOLOGICAL   ANATOMY. 

uterine  cavity,  incased  in  an  albuminous  fluid  and  an  outside 
fibrous  membrane  or  chorion  answering  to  the  egg-shell  in  the 
birds  (which  is  simply  calcified  chorion,  the  lime-salts  being 
deposited  through  the  structure,  while  the  vitelline  membrane 
(b),  zona  pelluclda,  inclosing  the  vitellus  (e)  or  yolk,  is  the 
same  in  both),  the  epithelium  of  the  chorion  begins  to  prolifi- 


Fig.  131. — A  Vertical  Section  from  the  tTterine  Mucous  Membrane,  showing  the  num- 
bers and  position  of  the  tubules.— Dalton.     a,  Free  surface  ;  b,  attached  surface. 


Fig.  132. — Same  tubules,  greatly  magnified. — Dalton. 

cate,  multiply  and  send  out  processes  similar  to  what  occurs  in 
the  zona  pelluclda  (a)  in  the  unimpregnated  ovum,  which 
rapidly  increase  in  size  in  the  rich  albuminous  fluid  poured 
out  by  the  uterine  follicles,  completely  surrounding  it,  uush- 
ing  the  decidual  membrane  before  it  and  filling  the  uterine 
cavity,  so  that  the  ovum  is  completely  submerged.  In  this 
thick  albuminous  substance,  containing  a  large  number  of  nu- 
cleated cells,  the  growing  tufts  of  the  chorion  are  embedded,  at 


PHYSIOLOGICAL   ANATOMY. 


323 


the  same  time  the  deeidua  reflexa,  formed  by  the  contiguous 
mucous  membrane,  grows  up  around  the  ovum,  bringing  the 
orifices  of  the  expanded  tubules  into  contact  with  the  project- 


Pig.  133.— Ovum  of  the  Rabbit,  from  a  Graafian  follicle  one-fiftieth  ot  an  Inch  in  diame- 


ter.—Waldeyer.  a,  Epithelium  of  the  ovum  ;  b,  vitelline  membrane,  showing 
the  radiating  striae  forming  the  pore  canals  through  which  the  spermatizoon  makes 
its  way  to  the  germinal  spot ;  c,  germinal  vesicle  ;  d,  germinal  spot ;  e,  vitellus. 


Fig.  134  —Impregnated  Uterus,  showing  connection  between  villosities  of  chorion  and 
decidual  membranes. — Dalton. 

ing  ends  of  the  growing  tufts,  which  readily  find  their  way  into 
the  lumen  of  the  expanded  uterine  tubules  (Fig.  134),  rapidly 
growing  and  branching  in  the  follicular  cavities,  which  increase 


324  PHYSIOLOGICAL    ANATOMY. 

correspondingly,  the  two  growing  pari  passu  with  each  other, 
the  branched  processes  constituting  the  villi  of  the  chorion 
(Fig.  135) ;  but  which  are  only  provisional,  the  greater  num- 
ber soon  fading  out,  so  as  to  make  the  chorion  "bald1'  save  in 
the  locality  elected  for  the  placenta,  the  portion  in  imme- 
diate contact  with  the  walls  of  the  womb,  where  they  rapidly 
increase  in  size,  as  also  the  uterine  follicles,  so  as  to  form  the 
placental  sinuses  ;  the  matter  being  one  of  continuous  growth 
and  amplification  on  the  part  of  both.  At  this  point  the 
capillary  loops  sent  into  the  villi  (they  had  previously  been 


Fig.  135. — Entire  Human  Ovum  of  Eighth  Week,  sixteen  lines  in  length  (not  reckoning 
the  tufts)  ;  the  surface  of  the  Chorion  partly  smooth,  and  partly  rendered  shaggy  by 
the  growth  of  tufts — Carpenter. 

only  amplified  epithelium,  inclosed  in  a  layer  of  basement 
membrane)  from  the  fcetal  vessels,  undergo  enlargement,  grow- 
ing rapidly,  while  the  uterine  sinuses  project  their  thin  endo- 
thelial lining  membrane  (extremely  thin)  into  the  placental 
sinuses,  forming  the  lining  membrane  to  them,  also,  whereby 
the  maternal  blood  is  brought  into  immediate  contact  with  tJi  <■ 
placental  villi,  for  producing  rapid  interchanges  between 
them,  while  special  arrangements  obtain  in  the  womb  itself 
(to  be  mentioned  presently)  for  rapidly  renewing  the  blood  in 
the  sinuses,  producing  a  continuous  stream  in  and  out  of  the 
sinuses,  which  the  scheme  calls  for. 

The  placenta,  then,  is  composed  of  both  fcetal  and  maternal 
structures  (Fig.  136).     The  structure  is  very  dense  ;   more  so 


PHYSIOLOGICAL    ANATOMY. 


325 


after  detacliment  than  before,  from  the  emptying  of  the  sinnses 
and  the  consequent  condensation  which  this  produces.  But 
the  placental  tufts,  with  the  intervening  boundary  wall,  com- 
posed of  follicular  membrane,  and  the  endothelial  layer  from 
the  uterine  sinuses  comes  out  sufficiently  clear  in  the  picture. 
The  extension  of  the  endothelial  lining  of  the  uterine  sinuses 
into  them,  so  as  to  bring  the  maternal  blood  into  immediate 
contact,  as  stated  by  Prof.  Dalton,  and  forcibly  illustrated  by 


Fig.  136. — Section  of  a  Fully  Formed  Placenta,  with  the  part  of  the  Uterus  to  which  it 
is  attached. — Cycl.  of  Anatomy,  Supplement,  1859;  Article,  "  Uterus  and  its 
Appendages,"  by  A.  Farre,  F.  R.  S.  /,  Umbilical  cord  ;  am,  amniotic  layer  ;  ch, 
chorion  ;  xis,  uterine  sinuses  ;  a,  a,  lines  indicating  the  point  where  the  placental  and 
uterine  structures  are  blended  ;  ate?,  where  the  structure  is  unusually  dense  and  the 
uterine  sinuses  are  small ;  dp,  showing  where  a  division-wall  between  the  placental 
tufts  is  divided  in  the  section  ;  S,  curling  arteries  of  uterus  :  v,  umbilical  vessels  ; 
vf,  branch  from  umbilical  vessels  to  the  membranes. 

the  following  diagram  (Fig.  137)  is  extremely  probable,  since 
it  is  precisely  what  is  needed  for  producing  free  action  in  the 
villi  for  pumping  the  fluids  into  thefcetal  circulation  and  ex- 
pelling waste  products.  Then,  again,  the  easy  manner  with 
which  the  placenta  is  peeled  off  and  detached  from  the  womb  is 
precisely  what  we  might  expect  from  this  mode  of  attachment, 
the  endothelial  lining  of  the  vessels  being  very  brittle  ;  so,  then, 
everything  is  in  correspondence.     But  were  the  uterine  arte- 


3JC  PHYSIOLOGICAL    ANATOMY. 

ries  and  veins  extended  into  the  placenta,  this  could  not  be 
done,  since  the  fibrous  and  muscular  structures  would  inhibit 
it,  and  parturition  would  have  a  Pandora' s  box  in  the  womb, 
leading   to  untold   evils,  and  in  all  probability  terminating 
fatally.     But  the  uterine  arteries  and  veins  terminate  by  ca- 
pillaries in  the  uterine  sinuses,  which  are  interspaces  or  canals 
formed  by   the  longitudinal  and  transverse  muscles   of  the 
womb,  and  lined  by  endothelium  extended  from  the  arterial 
and  venous  capillaries  so  as  to  form  a  common  reservoir  for  the 
vessels,  the  whole  bearing  close  resemblance  to  the  vascular 
spaces,  muscular  trabecule  and  terminal  vessels  in  the  corpora 
cavernosa  penis  (Fig.  204),  which  enables  the  rapid  filling  and 
emptying  of  the  hfemal  chambers  by  expanding  and  contracting 
the  muscles  and  the  vessels  by  the  action  of  the  special  nerv- 
ous forces  which  apply  in  the  organs  for  that  purpose,  press- 
ure applying  alike  to  both  organs,  and  under  the  pressure  in 
the    arterial    system,   the    reservoirs   are   rapidly    filled    in 
either  case,  and  as  readily  emptied  during  contraction.     It 
comes  to  this:    that   the  endothelial    lining  of    the  uterine 
sinuses   (Fig.    137,   c)  is  projected   into   the  vascular    inter- 
spaces in  the  placenta  forming  the  placental   sinuses,    and 
being  reflected  over  the  tufts,  these  are  inclosed  by  it  as  the 
fingers  by  a  glove,    every   one  receiving  an    investment  of 
uterine  endothelium,  so  that  the  maternal  blood  is  necessarily 
in  immediate  contact  with  them  ;  that  the  blood  is  readily  re- 
newed in  the  sinuses  commensurate  with  the  physiological 
requirements  in  the  embryo  ;    that  at  the  end  of  the  intra- 
uterine term  the  placenta  is  readily  detached  from  the  womb 
and  expelled  out  of  the  body  along  with  the  embryo,  the 
liquor  amnii  and  containing  membranes  under  the  action  of 
the  force  in  the  womb  and  walls  of  the  abdomen,  inclusive  of 
the  diaphragm,  all  the  parts  acting  in  concert  by  means  of  the 
correlation  of  the  nerves  in  the  medulla  oblongata,  or  the 
same  as  for  expelling  waste  products  in  the  rectum  and  blad- 
der, the  principle  being  the  same. 

And  by  looking  from  the  placental  sinuses  to  the  special  anat- 
omy in  the  placental  tufts  or  villi  (Fig.  138),  it  is  readily 
perceived  how  the  pumping  actions  may  be  set  up  in  them  for 
pumping  the  fluids  into  and  out  of  themselves,  the  same  as  in 


PHYSIOLOGICAL    ANATOMY. 


327 


the  lungs  when  the  animal  becomes  an  air-breather  at  the  end 
of  the  intra-uterine  term,  perfect  freedom  of  action  being 
alike  secured  in  both,  and  the  same  law  applying  for  produc- 
ing afflux  and  efflux  of  the  fluids  ;  but  a  still  more  close  re- 
semblance is  furnished  in  the  villi  of  the  intestinal  canal ; 
only  that  muscles  are  added  in  the  latter  for  producing 
more  energetic  action  and  a  larger  amount  of  work  to 
make  it  commensurate  with  the  force  expended  in  the  organ- 
ism.    Still,  it  is  manifest  from  the  special  anatomy  in  these 


Fig.  137. — Vertical  Section  of  Placenta,  showing  arrangement  of  maternal  and  fcetal 
vessels. — Dalton.    a,  a,  Chorion;  6,  6,  decidua  ;  c,  c,  c,  c,  orifices  of  uterine  sinuses 

organs,  that  absorption  must  go  on  energetically.  Thus, 
under  the  epithelium  {a,  a)  we  have  a  basis  of  connective  tissue 
fibres  (c,  c)  running  longitudinally  from  end  to  end  of  the 
organ,  and  by  expanding  and  contracting  a  pumping  action 
must  inevitably  be  produced  in  the  organ  for  aspirating 
and  propelling  the  fluids  to  and  from  the  foetus  and  the 
sinuses,  all  the  villi  acting  in  concert.  The  vascular  loops 
being  placed  within  the  band  of  connective-tissue  fibres  (b, 
c,  c),  must  necessarily  undergo  rhythmical  compression 
during  contraction,  thereby  increasing  pressure  in  them, 
causing  the  blood  to  flow  into  the  umbilical  vein  ;  while 
during  expansion  the  opposite  obtains,  causing  the  fluids  to 
flow  into  the  vessels  from  the  sinuses  and  umbilical  arteries 


328  PHYSIOLOGICAL    ANA'IOMV. 

from  a  reduction  in  pressure  which  this  effects,  while  the 
vessels,  by  acting  in  concert,  expanding  and  contracting  simul- 
taneously, should  increase  the  action  correspondingly.     Fur- 
thermore, it  is  easily  perceived  that  considerable  force    is 
necessary  for  compelling  the  fluids  through  the  animal  mem- 
branes, .since  this  outside  wall  of  the  placental  villus  (Fig. 
138)  is   composed  of  two  layers,    the  inner  one   of    epithe- 
lial cells  (a),  the  outer,  connective-tissue  fibres  (b)  ;    finally, 
the  capillary    wall   itself,    intervene   between   the    maternal 
and  total   blood,    and  that  interchange  would  have  to   be 
effected  through   these  membranes.       Now,   then,    for  com- 
pelling this  circumstance,  we  have  the  force  in  the  placental 
tupts  and  the  force  in  the  walls  of  the  womb  itself,  which  ex- 
pands and  contracts  regularly  and  rhythmically  for  diminish- 
ing   and    increasing  pressure    in   the  sinuses,    which    are 
simultaneous   with   the   movements    in    the    placenta,    only 
that   the  actions  alternate,    the   womb   contracting    as    the 
placenta  is  expanding,  and  vice  versa,    whereby   circulation 
in  the  uterine  and  placental  sinuses  is    readily  produced. 
In   other  words,   the   circulation  of   maternal  blood  in  the 
uterine  and  placental  sinuses  is  the  same  in  principle  as 
that  for  circulating  air  in  the  lungs — namely,  by  rhythmical 
changes  in  pressure  in  the  sinuses,  which  is  produced  by  the 
actions  taking  place  in  the  womb  and  placenta ;  taking  the 
placental  sinuses  to  represent  the  alveoli,  and  the  uterine  the 
tracheal  system,  the  blood  flowing  into  and  out  of  this  system 
of  canals  by  reflux  action  for  renewal,  just  as  the  pulmonic  air 
flows  into  and  out  of  the  tracheal  system  for  renewal,  only  that^ 
in  the  former  the  fluid  passes  into  the  venous  system  of  the 
mother  and  is  returned  by  the  arterial,  both  terminating  by 
capillary  openings  in  the  canals,  while  the  placental  souffle 
answers  to  the  respiratory  murmur.     And  when  the  womb 
contracts  for  increasing  pressure  in  the  sinuses,  the  action  is 
not  unlike  that  which  occurs  in  the  walls  of  the  intestines  for 
increasing  pressure  in  the  gut,  whereby  the  fluids  are  com- 
pelled more  rapidly  into  the  respective  villi,  flowing  from  high 
to  low  pressure.     Mirabile  ! 
The  following  facts  may  be  given  in  support  of  this  opinion : 
1.  The  womb  must  expand  and  contract  regularly  and  rhyth- 


PHYSIOLOGICAL    ANATOMY.  329 

mically  in  order  to  increase  circulation  in  the  sinuses,  since 
this  is  essential  for  changing  pressure  upon  the  blood,  while 
the  vast  number  of  muscles  and  nerves  in  the  walls  of  the 
womb  are  the  provision  for  more  energetic  action  than  is  possi- 
ble to  the  placenta,  which  is  composed  almost  entirely  of 
vascular  loops.' 

2.  The  womb  surrounds  the  embryo,  and  its  action  would 
therefore  be  more  effective  for  producing  the  changes  in  pres- 


Fig.  138. — Portion  of  one  of  the  Pcetal  Villi,  about  to  form  part  of  the  Placenta,  highly  * 
magnified. — Farre.  a,  a,  Its  cellular  covering  ;  b,  b,  b,  its  looped  vessels  ;  c,  c,  its  * 
basis  of  connective  tissue. 

sure  upon  it,  which  is  also  in  imitation  of  lohat  takes  place 
in  the  amnion  of  the  growing  chick,  it  being  rocked  to  and 
fro  in  the  egg  by  the  slow,  rhythmical  expansions  and  con- 
tractions taking  place  in  the  amnion,  and  by  alternating 
this  action  with  the  one  taking  place  in  the  placenta,  it  is 
readily  perceived  how  a  rapid  circulation  could  be  maintained 
in  the  maternal  and  placental  sinuses  for  compelling  corre- 
spondence between  this  and  the  energetic  circulation  in  the 
embryo,  since  it  all  forms  a  connected  whole.  This  would  also 
be  in  conformity  with  the  principle  in  the  circulation,   the 


330  PHYSIOLOGICAL    ANATOMY. 

blood  flowing  from  high  to  low  pressure.  As  the  placenta 
expands  for  reducing  pressure  within  itself,  and  for  increas- 
ing pressure  in  the  embryo,  simultaneously  and  pari  passu 
with  this  action,  the  entire  muscular  walls  of  the  womb  con- 
tract for  increasing  pressure  in  the  maternal  sinuses  for  com- 
pelling this  blood  in  the  placenta,  at  the  same  time  by  increas- 
ing pressure  upon  the  embryo  it  should  determine  a  more 
rapid  movement  from  the  latter  to  the  placenta ;  the  one  in- 
volving the  other.  But  when  the  placenta  contracts  for  in- 
creasing pressure  (thereby  compelling  the  blood  out  of  itself 
in  two  directions,  or  toward  the  embryo  and  the  maternal 
sinuses),  the  uterine  walls  expand  for  reducing  pressure  in 
the  embryo  and  the  uterine  sinuses,  thus  greatly  expediting 
the  placental  efflux ;  and,  taking  it  all  in  all,  there  can  be  very 
little  doubt  but  that  tine  force  which  is  represented  in  the 
muffled  murmur  of  the  placental  souffle  is  mainly  the  product, 
of  the  muscular  uterine  walls,  though  both  undoubtedly  con- 
tribute to  it. 

3.  This  action  in  the  uterus  and  placenta  would  account  for 
the  very  curious  and  suggestive  obliquity  in  that  portion  of 
the  canals  connecting  the  uterine  with  the  placental  sinuses, 
being  upon  a  line  almost  parallel  with  the  transverse  axis  of 
the  uterus  and  placenta,  which  is  precisely  what  is  called  for 
by  the  special  mechanics,  in  order  to  effect  the  lateral  or  to- 
and-fro  movements  which  the  separate  action  in  the  womb 
and  placenta  necessarily  involves,  the  vessels  simply  elongat- 
ing and  contracting  with  these,  as  the  case  may  be,  without 
interfering  with  the  calibre  of  the  tubes.  Were  the  vessels 
straight,  however,  it  would  be  utterly  impossible  to  operate 
this  mechanics,  since  the  lateral  movements  should  obliterate 
the  vessels  by  closing  the  calibre.  It  could  not  be  otherwise,  in 
the  very  nature  of  things.  In  the  diagram  (Fig.  137,  c),  the  ves- 
sels are  represented  as  nearly  perpendicular,  but  this  is  done 
simply  for  better  definition. 

4.  But  the  strongest  proof  of  this  higher  function  of 
the  womb  is  furnished  in  the  vascular  and  nervous  con- 
nections subsisting  between  it  and  the  maternal  organ- 
ism and  the  law  underlying  the  mechanics.  And  as  all 
tips  relates  to  circulation  for  building  up  and  elaborating 


PHYSIOLOGICAL    ANATOMY.  331 

the  embryo,  obviously  the  mechanics  for  increasing  circu- 
lation in  the  womb  commensurate  with  that  in  the  embryo 
should  extend  to  the  vessels  of  supply  or  the  feeders,  as  also 
the  discharging  vessels,  or  the  arteries  and  veins.  It  must  be 
shown  how  this  new  movement,  this  new  life  set  going  within 
the  other,  twines  its  arms  around  the  maternal  vessels  and 
feeds  itself  in  the  measure  of  its  necessities,  by  means  of  this 
pumping  action  in  the  womb  and  placenta  which  represents 
respiration.  The  spermatic  and  uterine  arteries  are  the  feeders 
(Fig.  139,  s,  u),  while  the  accompanying  veins  are  the  dis- 
charging vessels.  To  this  must,  be  added  the  uterine  lym- 
phatics, which  are  very  large  in  the  impregnated  womb.  They 
terminate  in  the  pelvic  and  lumbar  glands.  The  spermatic 
arteries  and  veins  have  similar  origin  and  termination,  as  in 
the  male,  while  the  uterine  artery  is  a  branch  of  the  internal 
iliac,  with  the  venous  return  through  a  vein  of  the  same  name. 
The  deeply  suggestive  fact  to  note  in  this  connection  is  that 
the  dense  plexuses  of  nerves  to  the  fundus  and  sides  of  the 
womb  converge  in  the  nervous  ganglia  about  these  vascular 
trunks,  or  the  spermatic  and  hypogastric  ganglia  (Fig.  140, 
€,  w,  r).  It  will  be  seen  that  the  nerves  to  the  fundus  (?;,  x) 
converge  in,  or  radiate  from,  the  spermatic  ganglion  (w)  which 
surrounds  the  spermatic  artery  and  vein  (e)  (which  corre- 
sponds with  the  attachment  of  the  placenta),  while  those  in 
the  neck  and  sides  from  the  hypogastric  ganglion  (r)  are 
brought  in  direct  relation  with  this  ganglion  by  means  of  in- 
tercommunicating nerves  (t)  for  unifying  the  action  through- 
out. 

Thus,  nervous  force  to  the  womb  is  literally  banked  upon 
the  Mood-vessels  ;  and  if  this  means  anything,  it  means  that 
circulation  shall  be  in  correspondence  with  the  physiological 
requirements,  or  supply  equal  to  demand,  the  cardinal  circum- 
stance being  the  growth  and  elaboration  of  the  embryo,  which 
is  the  object  and  purpose  of  the  organ,  the  others  being  sim- 
ply incidental. 

It  comes  to  this,  namely,  that  the  nervous  force  for  expand- 
ing and  contracting  the  uterine  sinuses  should  expand  and  con- 
tract the  uterine  blood-vessels  at  one  and  the  same  time,  there- 
by causing  afflux  and  efflux  of  blood  through  them  for  com- 


332 


PHYSIOLOGICAL    ANATOMY 


pelling  correspondence  throughout,  which  the  scheme  calls 
for.  Furthermore,  this  would  accord  with  the  action  in  the 
vessels  in  respiration,  as  indicated  by  the  undulations  in 
arterial  tension.  But  would  not  this  mechanics  interfere  with 
the  due  circulation  of  blood  in  the  placental  sinuses  I  Cer- 
tainly not,  and  for  the  following  reasons  : 


Fig.  139.— Showing  the  Arteries  and  Veins  to  the  Womb  (left  side).  S,  spermatic  artery 
and  veins  (ovarian)  ;  U,  uterine  artery  and  vein  ;  1,  vessels  passing  between  the 
muscular  fibres ;  2,  peritonaeum  ;  G,  Fallopian  tube  ;  F,  ligament  of  ovary  ;  i?, 
round  ligament  ;  H,  inferior  ligament  or  duplicature  of  peritonaeum  corresponding 
with  Douglas'  cul  de  sac  ;  V,  vagina. 

1.  Efflux  of  blood  in  the  uterine  sinuses  through  the  veins 
is  by  means  of  capillary  vessels,  which  tend  to  retard  escape, 
while  the  flow  of  blood  into  and  out  of  the  placental  sinuses 
takes  place  through  large  canals  (Fig.  137,  c,  c),  whereby  afflux 
and  efflux  has  corresponding  increase.  But  in  addition  to  this, 
the  expansion  in  the  placenta,  which  occurs  during  systole  of 


NERVES   TO   THE   GRAVID   WOMB. 


'333 


A,  the  fundus  and  body  of  the  uterus,  having  the  peritoneum 
di^secred  off    from  left    side ;    B,  the  vagina  covered  with 
nerves  proceeding  from  the  inferior  b^rd-r  of  the  left  hypo- 
gastric ganglion  ;    C,   rectum  ; 
£>,  left  ovarium  and  Fallopian, 
tube  ;    E,  trunk  of  left   sper- 
matic vein  and  artery 


Fig.  140.— Ganglia  and  Nerves  of  the     entnfflw    feft^hvpoVastYicT  ganglia,  "and    giving    oft 

m         -j  ttj-  ,    ,,      -n,     ■,      c  ,.        hrMiiphps  to  the  left  pub-peritoneal  ganglia;  P. nsemorr- 

Gravid  Uterus  at  the  End  of  the     hoWal  nerves  and  art.rv  r  <?,  sacral  nerves  entering  the 

Ninth  Month  — R    Lee  «h  'le  cuter  surface  of  the  hypogastric  ganelion ;  iJ,  left. 

INintn  Jiiontn.     it.  l^ee.  hypogastric  ganglion  with  the  arteries  '"Jerted^nerveg 

of  the  vagina ;    T,  nerves  with  an  injected  artery  proceeding  from  the  upper  part  ot  Hit  hypogasn  u, 

ganglion  along  the  body  of  the  uterus,  and  terminating    n  the  left  spermatic  ganglion  ,   <^^x"'"il" 

tion  of  these  nerves  and  the  branches  which  they  gi>-e  off  to  the  sub-peritoneal  plexuvte.i^some, 

nerves  passing  upward  beneath  th    sub-peritoneal  plexuses,  and  anastomoses  treely  wmi  tnem  ,       , 

left  spermatic  ganglion,  in  v-hich  the  nerves  and  artery  from  the   hypogastric  ganglion,  ana  me> 

branches  of  the  left  sub-peritoneal  plexuses  terminate  ,and  from  wh'  h  The  nerves  of  the  ru°Q;'.°  }"J  {, 

are  supplied  ;  X,  left  sub-peritoneal  plexuses  covering  the  body  of  the  uterus;    I.leftsu^b^eHtoneaL 

ganglion,  with  numerous  branches  of  nerves  extending  between  it  and  left  lipoipstnc  nene  dnu 

ganglia  ;  Z,  left  common  iliac  artery  cut  across  and  turned  aside  to  expose  left  hypogastr  e  nerve  ami. 

ganglion. 


334  THE   ACTION   IN  THE   MATEKNAL   VESSELS. 

the  womb,  should  also  determine  the  blood  in  this  direction, 
so  that  most  ample  arrangements  exist  for  producing  the  freest 
circulation  in  the  placental  sinuses. 

2.  During  diastole  in  the  womb  and  its  sinuses,  for  aspirat- 
ing the  venous  blood  in  the  placental  sinuses,  the  high 
pressure  in  the  arterial  system  causes  the  arterial  blood  to 
flow  into  the  uterine  sinuses  as  rapidly  as  the  blood  coming 
from  the  placenta  ;  at  the  same  time,  the  valves  in  the  veins 
of  the  uterus  obviate  reflux  in  the  venous  system.  And  when 
the  womb  contracts  for  compelling  the  blood  in  the  uterine 
sinuses  into  the  placental  sinuses  and  venous  system,  the 
pressure  in  the  arterial  system  inhibits  reflux  in  this 
direction.  Thus  the  mechanics  for  increasing  circulation 
in  the  uterine  and  placental  sinuses  is  complete  in  every 
respect ;  nor  does  it  require  extensive  expansion  and  con- 
traction in  the  womb  in  order  to  effect  increased  circu- 
lation through  the  sinuses,  but  a  moderate  and  limited 
amount,  sufficient  only  for  producing  rhythmical  changes  in 
pressure  upon  the  blood,  as  must  appear  obvious  ;  something 
similar  to  what  takes  place  in  the  spongiae,  which  have  muscles 
(Norway)  for  producing  a  more  rapid  circulation  in  and  out 
of  this  canal  system  ;  while  in  reference  to  the  action  in  the 
vessels,  their  rhythmical  expansions  and  contractions  with  the 
movements  in  the  womb  and  placenta,  this  has  its  analogue  in 
the  mammalia,  the  maternal  arterial  system,  which  maintains  a 
similar  action  synchronous  with  respiration,  as  has  been  fully 
shown  ;  and  is  it  unreasonable  to  make  similar  deduction  for 
the  womb  and  embryo  for  increasing  circulation  in  them  as 
well,  more  especially  when  no  other  means  exist  for  increasing 
circulation  and  for  making  it  commensurate  with  the  physio- 
logical requirements  in  the  growing  embryo  ?     I  think  not. 

This  expansion  of  the  womb  and  arterial  feeders  would  ex- 
plain the  sudden  and  enormous  escape  of  arterial  blood  in  post- 
partem  haemorrhage,  for  the  placenta  being  no  longer  attached, 
the  flow  of  blood  through  the  uterine  sinuses  must  necessarily 
be  purely  arterial.  In  other  words,  when  contraction  ceases 
and  the  movement  of  expansion  sets  in,  the  flood-gates  are 
thrown  open  to  the  arterial  system,  hence  the  rush  of  blood 
from  it. 


1HE   KHYTHMIC    CENTEE   FOR   THE   WOMB.  335 

Inreference  to  the  nervous  centre  for  this  pumping  action  in 
tlie  womb,  which  answers  to  respiration  in  the  fcetus.  There 
can  be  very  little  doubt  but  that  the  spinal  cord  functions  as 
the  common  reflex  centre  of  nervous  force  for  producing  the 
rhythmical  expansions  and  contractions  in  the  gravid  womb 
simulating  respiration,  in  short,  that  the  lumbar  enlarge- 
ment of  the  spinal  medulla  is  in  point  of  fact  the  respiratory 
centre  for  the  foetus,  regarding  the  pumping  action  in  the 
womb  and  placenta  as  the  respiratory  movement,  of  which 
there  can  be  but  little  doubt.  And  since  the  local  nervous 
.ganglia  control  the  local  actions,  maintaining  them  also  in 
action  as  in  the  case  of  the  locomotor  apparatus,  is  it  illogical 
to  make  the  same  deduction  for  the  womb  and  the  reflux  cen- 
tre in  the  spinal  cord  ?  I  think  not.  In  short,  there  must  be 
a  nervous  centre  for  coordinating  and  unifying  the  movements 
in  every  organ,  in  the  uterus  as  well  as  elsewhere,  as  must 
appear  obvious.  The  following  are  the  reasons  for  this  allega- 
tion : 

1.  The  very  intimate  connection  subsisting  between  the 
womb  and  spinal  cord  by  means  of  the  hypogastric  ganglia  and 
sacral  plexus  with  the  intercommunicating  nerves  extending 
from  the  spinal  medulla  (Fig.  116). 

y.  The  fact  that  the  lumbar  portion  of  the  spinal  cord  under- 
goes enlargement  during  gestation. 

3.  The  fact  that  reflex  action  in  the  womb  is  readily  pro- 
duced by  applications  to  the  slcin  surface,  a  circumstance 
well  known  and  practiced  by  the  profession.  All  these 
facts,  then,  fall  readily  into  line  when  viewed  from  this 
stand-point ;  and,  no  scientific  reason  existing  to  the  con- 
trary, we  may  conclude  the  fact  as  logically  proven,  while 
the  vast  number  of  the  muscles  and  nerves  in  the  womb 
make  it  absolutely  certain  there  is  such  rhythmic  movement 
going  on.  Indeed,  the  gist  of  the  question  is  not  whether  there 
is  a  reflex  centre  in  the  spinal  cord  for  the  womb,  but  whether 
this  centre  produces  the  rhythmical  expansions  and  contrac- 
tions in  that  organ,  as  alleged.  A  full  and  sufficient  answer  to 
which  is  furnished  in  the  fact  of  the  total  absorption  of  the 
relative  phenomena,  anatomical  and  physiological,  otherwise 
utterly  inexplicable  ;  while  underneath  all  which  is  the  organic 


330  THE    RHYTHMIC    CEWTEE   FOR   THE   WOMB. 

law  on  which  animal  life  itself  is  constructed,  calling  for  rhyth- 
mical changes  in  pressure  in  the  contents  of  the  gravid  womb 
and  in  the  uterine  sinuses  for  increasing  circulation,  and  for 
making  it  commensurate  with  the  physiological  requirements 
in  the  embryo ;  otherwise  impossible.  Nor  is  it  reasonable 
that  the  enormous  number  of  muscles  and  nerves  in  the  womb 
are  for  compelling  out  the  contents  at  the  end  of  the  term  sim- 
ply ;  but,  on  the  contrary,  tluit  they  perform  an  active  and 
essential  part  in  the  worlc  of  construction  which  precedes  ex- 
pulsion. Beyond  a  shadow  of  a  doubt,  they  are  not  idle  in  all 
this  while,  especially  when  supreme  necessity  would  have  it 
otherwise  ;  on  the  contrary,  they  are  evolved  as  force  is  needed 
for  carrying  on  circulation,  while  at  the  end  of  the  term  they 
are  available  for  assisting  in  expelling  the  embryo  ;  hence,  per- 
form an  active  role  from  the  beginning  to  the  end  of  their  ex- 
istence, as  is  ever  the  case  with  the  muscles  and  nerves. 

One  other  circumstance  in  this  connection,  namely :  the 
very  tortuous  course  of  the  arteries  in  the  womb  (Fig.  139,  u), 
which  undoubtedly  has  reference  to  this  action  in  the  womb, 
permitting  the  rhythmical  expansions  and  contractions  to 
take  place  without  involving  strain  to  the  vessels,  other- 
wise inevitable.  The  veins,  it  will  be  perceived,  take  a 
straight  course,  while  the  arteries  are  serpentine  or  bent 
upon  themselves.  Ihis  is  due  to  the  fact  that  the  veins 
are  more  extensile,  and  possess  greater  powers  of  elongating 
and  shortening  than  the  arteries,  the  yellow  elastic  coat  of 
the  latter  tending  to  limit  their  actions.  It  will  be  remem- 
bered that  this  circumstance  has  forcible  illustration  in  the 
splenic  artery  and  vein,  the  former  being  almos-t  twice  the 
length  of  the  latter,  to  allow  for  expansion  in  the  stomach 
when  food  is  taken  ;  otherwise,  this  would  involve  rjrodigious 
strain  to  the  vessels,  with  great  reduction  of  the  calibre.  But 
the  same  remark  will  apply  to  the  vessels  of  all  the  hollow 
viscera.  Thus,  everything  is  in  correspondence.  Of  course, 
the  movement  in  the  womb  and  placenta  is  necessarily  more 
limited  than  in  the  lungs,  in  which  considerable  space  is  re- 
quired for  sucking  in  the  air  simultaneously  with  the  venous 
blood,  but  which  would  not  apply  for  the  fcetal  circulation,  as 
the  oxygen  is  furnished  by  arterial  blood  at  one  and  the  same 


EXPELLING   THE    UTERINE  CONTENTS.  337 

time  with,  the  nutritive  and  force-producing  elements,  which 
the  scheme  calls  for  in  order  to  generate  force,  since  it  is  by  a 
combination  of  the  two  that  force  is  evolved,  as  before  re- 
marked. In  fine,  differentiation  in  the  organs  cannot,  for 
obvious  reasons,  work  any  change  in  the  fundamental  principle 
underlying  the  mechanics  for  increasing  circulation,  which  is 
by  rhythmical  changes  in  pressure  involving  a  pumping  action 
for  compelling  the  commerce  in  the  blood-vessels  and  expelling 
waste  products,  while  the  speed  of  the  currents  thus  produced 
is  determined  by  the  rapidity  and  energy  of  the  rhythmical 
expansions  and  contractions  pervading  the  organs,  inclusive  of 
the  heart  and  vessels,  since  it  all  forms  a  connected  movement 
for  increasing  circulation  between  the  cell-brood  and  environ- 
ment, from  which  everything  is  obtained  and  into  which, 
in  due  time,  everything  is  returned  for  redistribution,  in  the 
embryo  the  same  as  in  the  maternal  tissues,  only  the  journey 
to  and  from  the  environment  is  by  way  of  the  maternal  blood- 
vessels, inclusive  of  the  uterine  sinuses,  the  common  ground 
where  interchange  is  effected  betwen  the  maternal  and  foetal 
blood. 

With  the  expiration  of  the  intra-uterine  term,  expansion  of 
the  maternal  passages  sets  in  for  reducing  resistance  to  the 
egress  of  the  embryo,  and  the  womb  and  abdomen  contracting 
simultaneously  for  increasing  pressure  in  the  womb,  the  con- 
tents are  compelled  into  the  environment.  Here,  as  else- 
where, the  law  of  pressure  applies  for  compelling  movement 
in  the  contents  ofthehollow  viscera,  for  which  special  adjust- 
ments obtain  in  the  organs  and  organism,  the  underlying 
principle  being  ryhthmical  changes  in  pressure. 

Pressure  being  invisible,  it  is  difficult  to  realize  the  impor- 
tant relations  it  sustains  to  the  mechanics  and  the  enormous 
role  it  performs  in  the  organism  ;  nevertheless,  the  fact  is  in- 
controvertible that  from  centre  to  circumference,  and  from 
surface  to  surface  of  the  body,  it  is  the  fundamental  and  con- 
trolling circumstance,  the  foundation,  so  to  speak,  of  the  tem- 
ple, pervading  the  superstructure,  and  interwoven  with  all  the 
phenomena,  which  spring  out  of  it  as  waters  from  a  fountain. 

It  is  passing  strange  the  matter  should  have  escaped  atten- 
tion so  long,  especially  in  this  age,  when  thought  is  reaching 


338     CHANGES    IN   THE   MECHANICS    PRODUCED    BY   BIRTH. 

down  into  the  organic  basis  of  life.  Indeed,  one  needs  to  go 
there  if  he  would  unravel  the  tangled  skein  in  animal  struc- 
ture and  function,  since  the  definite  arrangements  that  obtain 
in  the  organs  with  every  stage  in  development  show  unmis- 
takably a  common  relation  to  fundamental  forces  in  Nature 
underlying  it  all,  notably  pressure  and  gravitation,  while  the 
arrangements  which  obtain  in  the  structures  represent  the 
relative  adjustments  for  special  work,  and  in  the  mea  ure 
of  it. 

Respiration  in  the  New-born  :  The  Change  in  Mechanics 
which  this  Involves. — The  first  thing  in  the  new-born  is  to  start 
respiration  for  compelling  in  the  commerce  in  the  environ- 
ment in  place  of  the  uterine  sinuses,  and  the  action  in  the 
placenta  for  which  this  is  the  substitute,  the  oxygen  pass- 
ing in  by  way  of  the  lungs  and  the  aliment  through  the 
intestinal  canal.  But  it  requires  fresh  adjustments  in  the 
mechanics  of  circulation  to  bring  it  in  correspondence  with 
this  circumstance ;  notably,  circulation  of  the  blood  in  the 
lungs,  and  the  attaching  of  the  intestinal  apparatus  to  this 
movement  by  means  of  the  nerves  connecting  in  the  medulla 
oblongata,  a  matter  which  has  already  been  fully  considered 
in  the  preceding  pages.  The  first  thing,  therefore,  is  to  start 
respiration,  when  it  will  be  in  order  to  consider  how  the  me- 
chanics in  circulation  swings  into  this  pendulum  movement  for 
compelling  correspondence  throughout,  with  the  blood  ever 
flowing  from  high  to  low  pressure,  in  conformity  with  the  or- 
ganic law  underlying  the  organism  itself.  One  end  of  the 
nervous  system,  so  to  speak,  is  spread  out  in  the  skin  surface, 
the  other  through  the  organism,  while  the  medulla  oblongata 
functions  as  the  common  centre  to  it  all ;  any  impression, 
therefore,  made  upon  the  skin  surface  is  promptly  reflected  to 
the  medulla  oblongata,  thence  over  all  the  structures  for  pro- 
ducing the  reflex  actions  connected  with  respiration  and  circu- 
lation. The  irritations  attendant  upon  parturition  from  fric- 
tion against  the  maternal  structures  are  calculated  to  produce 
these  reflex  actions  ;  but  the  contact  of  the  sentient  surface 
with  the  stimulus  in  the  atmosphere  itself  would  also  excite 
it.  And  if  the  child  should  be  injured  by  the  rude  experi- 
ences incidental  to  parturition,  a  yet  more  powerful  means  for 


CHANGES   IN   THE   MECHANICS   PRODUCED    BY   BIRTH.     339 

exciting  the  reflex  actions  connected  with  respiration  is  fur- 
nished by  the  sudden  application  of  cold  to  the  surface,  as 
in  sprinkling  cold  water  upon  it,*  or  a  sudden,  sharp  slap 
with  the  open  hand  may  be  substituted  instead,  as  is  com- 
monly practiced.  Last,  but  not  least,  carbonic  acid,  as  it  ac- 
cumulates in  the  blood,  acts  as  a  special  stimulus  to  respira- 
tion. It  cries  out  in  pain,  and,  presto  !  the  haemal  mechanics 
is  changed.  The  low  pressure  which  is  produced  in  the  alveoli 
by  expansion  of  the  lungs  during  inspiration  compels  simul- 
taneous afflux  of  air  and  blood  in  the  alveoli  ;  while  the  high 
pressure  which  is  produced  by  the  subsequent  contraction 
during  expiration  causes  simultaneous  efflux  in  these  fluids, 
which  flow  from  high  to  low  pressure  in  conformity  with 
organic  law — the  one  flowing  out  by  reflux  action  through  the 
route  of  ingress,  the  other  passing  into  the  left  chambers  of 
the  heart  and  arterial  system  on  its  way  to  the  cell-brood,  as 
has  already  been  described  in  the  air-breather.  This  aban- 
donment of  the  old  route  for  the  new  is  readily  explained, 
since  it  is  in  strict  accordance  with  physical  law,  being  in  the 
direction  of  least  resistance. 

For  example,  we  begin  the  mechanics  with  high  rjressure  in 
the  arterial  system,  since  this  extends  through  the  ductus  ar- 
teriosus to  the  semilunar  valves  of  the  pulmonary  artery, 
the  floor  of  support  to  the  arterial  column.  Hence,  when  the 
alveoli  expand  during  inspiration  for  sucking  in  air  through 
the  trachea,  the  high  pressure  in  the  pulmonary  artery  and 
ductus  arteriosus  compels  this  blood  to  flow  straight  on  to  the 
low-pressure  areas  in  the  alveoli  simultaneously  with  the 
afflux  of  air,  or  in  the  direction  of  least  resistance,  in  place  of 
forcing  its  way  into  the  arterial  system  against  high  pressure, 
which  would  be  contrary  to  law.  And  the  ductus  arteriosus, 
though  still  filled  with  blood,  as  in  the  case  of  an  artery,  be- 
yond the  ligature  to  where  a  collateral  branch  is  given  off, 
shrinks  and  contracts  till  it  becomes  a  solid,  impervioas  cord. 

For  closing  the  foramen    ovale,  the  following  mechanics 

*  The  intimate  connection  subsisting  between  the  respiratory  cpntre  and 
the  skin  surface  is  of  easy  demonstration  in  the  adult  by  the  same  means. 
For  example,  every  impact  of  cold  water  against  the  skin  products  spasmodic 
inspiration  or  expansion  in  the  lungs  ;  not  deep,  however,  but  very  energetic. 


340     CHANGES   IN    'JHE   MECHANICS    PRODUCED    BY    BIRTH. 

apply :  After  birth,  the  inpour  of  blood  in  the  left  auricle 
by  way  of  the  pulmonary  veins  is  as  rapid  as  it  is  in  the 
right  auricle  through  the  venae  cavae,  and  with  pressure  at 
equilibrium  in  the  two  auricles,  this  at  once  suspends  all 
tendency  in  the  blood  to  pass  from  one  side  into  the  other 
during  auricular  diastole ;  while  during  the  auricular  systole 
and  the  high  pressure  this  produces  in  the  auricles,  causes 
the  blood  to  flow  into  the  expanding  ventricles,  where  low 
pressure  invites  it,  at  the  same  time  gravitation  also  should 
compel  it  in  this  direction,  since  the  ventricles  are  under  the 
auricles,  the  same  applying  for  either  auricle.  Thus,  a  dual 
force  applies  (suction  and  gravitation)  for  compelling  this 
blood  into  the  ventricles  during  the  auricular  systole,  and 
the  foramen  ovale,  being  thus  abandoned,  is  closed  and 
obliterated  by  membranous  formation. 

But  in  intra-uterine  life  the  matter  is  different ;  here  the 
wThole  blood  is  poured  into  the  right  auricle,  that  from  the 
upper  cava  passing  at  once  into  the  right  ventricle,  while  that 
in  the  lower  cava  (which  includes  the  blood  from  the  umbilical 
vein)  passes  through  the  right  into  the  left  auricle,  with  which 
it  directly  communicates,  guided  by  the  Eustachian  valve,  but 
also  pushed  over  and  deflected  in  this  direction  by  the  weight 
of  the  descending  current  from  the  npper  cava ;  but  if  the 
head  be  downward  (which  is  generally  the  case),  then  by  its  own 
weight  the  blood  would  gravitate  in  this  direction,  the  influx 
of  blood  from  the  upper  cava  also  compelling  it.  And  with 
the  absence  of  blood  as  a  counter-force  in  the  left  auricle,  this 
blood  is  necessarily  compelled  into  the  latter,  thence  into  the 
left  ventricle  and  aorta,  while  that  in  the  right  ventricle  passes 
into  the  arterial  system  at  the  aortic  arch  by  way  of  the  pulmo- 
nary artery  and  ductus  arteriosus.  After  birth,  however,  the 
pumping  action  in  the  lungs  reverses  all  this,  in  maimer  as 
above  described.  The  pulmonary  artery  in  the  embryo,  in 
place  of  discharging  through  the  lungs,  left  auricle  and  ven- 
tricle, empties  its  blood  at  once  into  the  aorta  as  it  passes 
under  the  arch,  and  which  is  also  in  the  direction  of  least  re- 
sistance, ^ince  it  is  impossible  for  this  blood  to  thread  its  way 
through  the  capillary  meshes  of  the  unexpanded  alveoli  ;  a  cir- 
cumstance which  has  forcible  illustration  in  the  air-breather, 


PHYSIOLOGY   OF    THE  ALR-<  HAMBER    IN   THE  EGG.         341 

and  when  the  alveoli  are  filled  with  residual  air,  by  simply 
inhibiting  inspiration  by  closing  the  mouth  and  nose  so  as  to 
prevent  expansion  in  the  lungs,  the  blood,  in  consequence, 
rapidly  accumulating  in  the  right  side  of  the  heart  and  venous 
system.  In  the  space  of  a  minute  there  is  lividity  of  the  lips 
and  whole  cutaneous  surface  from  venous  stasis  in  the  sys- 
temic capillaries.  If  longer  than  this,  an  appalling  venous 
suffusion  pervades  the  surface ;  in  the  face  most,  for  this  is 
the  most  vascular  portion,  with  the  large  venous  trunks  in 
close  proximity  to  the  heart.  Even  the  eyes  are  forced  out- 
ward, becoming  prominent  from  distension  of  the  intra-orbital 
veins  caused  by  obstruction  in  the  cavernous  and  lateral 
sinuses  But  the  instant  the  obstruction  is  removed  and  the 
lungs  are  permitted  to  expand,  so  as  to  reduce  the  intra- 
pulmonic  pressure,  the  dammed-up  blood  surges  into  the 
alveoli,  and  all  runs  on  as  before.  In  other  words,  the  heart 
and  vessels  are  unable  to  carry  on  circulation  in  the  absence  of 
the  pumping  action  in  the  lungs,  for  which  afflux  and  efflux 
of  air  is  essential ;  all  of  which  has  been  sufficiently  explained 
in  the  preceding  pages. 

Concerning  Incubation  and  Circulation  in  the  Egg. — Why 
should  there  be  an  air-chamber  to  the  egg  (Figs.  141  and  142)  ? 
We  are  now  prepared  to  furnish  a  scientific  explanation  to  this 
physiological  problem,  otherwise  inexplicable,  viz. :  the  con- 
tents of  the  egg  for  developing  the  chick  are  inclosed  by  a  firm, 
unyielding  wall  of  living  marble,  and  since  the  animal  circula- 
tion is  dependent  upon  rapid  rhythmical  changes  in  pressure, 
it  follows  that  provision  should  be  made  within  the  shell  for 
effecting  this  ;  otherwise  the  actions  in  the  heart  and  vessels 
could  not  take  place.  This  air-chamber  (a),  together  with  the 
important  relations  it  sustains  to  circulation  and  elaboration  in 
the  growing  chick,  organilogically,  therefore,  must  be  regarded 
as  one  of  the  most  essential  and  important  elements  in  egg- 
structure,  the  underlying  principle  to  all  the  nutritive  changes 
which  are  ushered  in  under  the  action  of  external  tempera- 
ture. The  accompanying  illustration  (Fig.  142)  will  serve  for 
impressing  the  matter. 

The  discipline  in  the  nutritive  processes  requires  the  blood 
to  be  brought  from  the  vitellus  and  aerated  in  the  allantois, 


342 


PHYSIOLOGY   OF    THE  AIR-CHAMBER    IN  THE  EGG. 


thence  to  be  dispatched  through  the  body  territories.  Accord- 
ingly, two  great  venous  trunks  (omphalo-meseraic  veins),  one 
in  each  fold  of  the  splanchnopleure,  embracing  the  vitellus, 
are  the  first  evolved,  while  at  the  terminal  ends  or  confluence 
the  heart  is  formed  by  the  blending  of  the  walls  of  these  venous 
trunks. 


Fig.  141. — Anatomy  of  the  Egg. — Jones,  a,  b.  Air- vesicle  ;  6,  arrow  indicating  the 
position  of  the  central  axis  of  the  egg  ;  c,  the  yolk  ;  /.  Purkinjean  vesicle  ;  g, 
cicatricula  ;  h,  thickening  of  the  vitelline  membrane  ;  e,  canal  leading  to  d,  the  cen- 
tral chamber  of  the  yolk. 


Fig.  142. — Egg  of  Fowl  in  Process  of  Development,  showing  area  vasculosa,  with  vitelline 
circulation,  terminal  sinus,  etc. — Dalton. 

According  to  His.  the  heart  is  developed  by  the  coalescence 
of  a  layer  of  the  splanchnopleure  with  a  similar  layer  from  the 
somatopleure,  the  hollow  cavity  formed  by  the  union  being  in 
free  communication  with  the  adjacent  omphalo-meseraic  veins. 
According  to  Foster  and  Balfour,  "  the  upper  end  of  the  heart 
is  developed  out  of  the  mesoblast  of  the  splanchnopleure,"  but 
"increases  in  length  step  by  step  at  the  expense  of  the  con- 


PHYSIOLOGY    OF   THE    CIKCULATIOX.  B43 

tinually  coalescing  omphalo-meseraic  veins."  Thus  the  fact  is 
undeniable,  that  the  heart  is  developed  in  the  venous  system 
in  connection  with  the  vitellus.  The  rhythmical  expansions 
and  contractions  in  this  organ  (the  punctum  saliens  of  early 
authors)  serve  to  pump  the  vitelline  fluids  in  the  bulbus 
arteriosus  and  the  two  aortse  which  are  developing  at  the  other 
end  of  the  heart.  But  for  this  air-cushion  within  the  egg  (Fig. 
141,  a,  b),  neither  these  rhythmical  expansions  in  the  heart 
nor  the  changes  in  pressure  for  compelling  circulation  in  the 
vitelline  fluids  could  take  place,  since  the  unyielding  shell 
would  inhibit  these  actions,  as  must  appear  obvious. 

Respiration  is  provided  for  in  the  following  manner  :  The 
allantois  (a  diverticulum  of  the  intestinal  canal)  is  pushed  out 
around  the  amnion  which  contains  the  embryo,  and  expanding 
its  vast  capillary  network  of  vessels  (whose  footstalks  spring 
from  the  two  iliac  arteries,  as  do  the  umbilical  arteries  in  the 
mammalian  embryo)  against  the  shell-membrane  or  chorion, 
becomes  the  respiratory  organ  of  the  chick,  by  means  of  which 
the  venous  blood  is  constantly  arterialized,  the  oxygen  passing 
in  and  the  carbonic  acid  passing  out  through  the  pores  in  the 
shell  by  the  action  of  the  polar  forces.  But  "at  the  time  the 
heart  first  begins  to  beat,  the  capillary  system  of  the  vascular 
and  pellucid  areas  is  not  yet  completed,  and  the  fluid  which  is 
at  first  driven  by  the  heart  contains,  according  to  most  ob- 
servers, very  few  corpuscles The  course  of  the 

blood  then,  during  the  latter  half  of  the  second  day,  may  be 
described  as  follows :  The  blood  brought  by  the  omphalo- 
meseraic  veins  falls  into  the  twisted  cavity  of  the  heart,  and  is 
driven  thence  through  the  bulbus  arteriosus  and  aortic  arches 
into  the  aortic  trunk.  From  the  aorta  by  far  the  greater  part 
of  the  blood  flows  into  the  omphalo-meseraic  arteries,  only  a 
small  amount  passing  on  into  the  caudal  terminations.  From 
the  capillary  network  of  the  vascular  and  pellucid  area  into 
which  the  omphalo-meseraic  arteries  discharge  their  contents, 
part  of  the  blood  is  gathered  up  at  once  into  the  lateral  or 
direct  trunks  of  the  omphalo-meseraic  veins.  Part,  however, 
goes  into  the  middle  region  of  each  lateral  half  of  the  sinus 
terminalis,  and  there  divides  on  each  side  into  two  streams. 
One  stream  and  that  the  larger  one,  flows  in  a  forward  direc- 


344  PHYSIOLOGY    OF  THE   CIRCULATION. 

tion  until  it  reaches  the  point  opposite  the  head,  thence  it 
returns  by  the  veins  spoken  ol  above,  straight  to  the  omphalo- 
meseraic  trunks.  The  other  stream  flows  backward,  and  be- 
comes lost  at  the  point  opposite  the  tail."* 

The  following  from  the  same  authors  is  deeply  suggestive : 
"  ISoon  after  its  formation  the  heart  begins  to  beat,  at  first  slow 
and  rare  pulsations,  beginning  at  the  venous  and  passing  on  to 
the  arterial  end.  It  is  of  some  interest  to  note  that  its  func- 
tional activity  commences  long  before  the  cells  of  which  it  is 
composed  shoio  any  distinct  differentiation  into  muscular  or 
nervous  elements."  It  would  be  difficult  to  overestimate  this 
circumstance,  since  it  establishes  beyond  peradventure  the 
power  in  the  higher  as  in  lower  animals  to  effect  rhythmical 
expansions  and  contractions  in  the  soft  tissues  in  the  absence 
of  any  muscle  or  nerve  for  producing  them.  But  at  present 
the  significance  of  these  rhythmical  expansions  and  contractions 
taking  place  in  the  heart  concerns  us  most,  since  the  manifest 
purpose  is  to  increase  circulation  ;  and  as  this  can  only  be  done 
by  pumping  the  blood  of  the  omphalo-meseraic  veins,  it  fol- 
lows that  both  expansion  and  contraction  is  necessary  for 
accomplishing  this — the  one  for  aspirating,  the  other  for  pro- 
pelling it.  This  would  explain  why  the  action  should  begin 
at  this  end  of  the  heart.  Of  course,  as  the  area  of  circu- 
lation increases  this  would  call  for  corresponding  increase  of 
force  for  effecting  it.  Accordingly,  pressure  is  increased  at 
the  same  time  that  additional  force  is  placed  upon  it ;  notably, 
by  the  amniotic  fluid  and  by  the  action  in  the  amnion,  the 
muscles  and  nerves  in  the  heart  and  vessels  being  in  cor- 
respondence. 

The  amnion  closes  around  the  embryo  of  the  chick  on  the 
fourth  day,  and  on  the  fifth  fluid  begins  to  collect  in  the  sac, 
and  by  the  seventh  the  embryo  is  submerged  in  a  considerable 
quantity  of  water.  "By  the  seventh  day  very  obvious  move- 
ments begin  to  appear  in  the  amnion  itself  ;  slow  vermicular 
contractions  creep  rhythmically  over  it.  The  amnion,  in  fact, 
begins  to  pulsate  slowly  and  rhythmically,  and  by  its  pulsa- 
tions the  embryo  is  rocked  to  and  fro  in  the  egg.  This  pulsation 

*  Foster  and  Balfour's  "Embryology." 


PHYSIOLOGY   OF   THE   CIRCULATION.  345 

is  due,  probably,  to  the  contraction  of  involuntary  muscular 
fibres,  which  seem  to  be  present  in  the  attenuated  portion  of 
the  mesoblast,  forming  part  of  the  amniotic  fold"  (Foster  and 
Balfour). 

The  physiological  significance  Of  this  accumulation  of  amni- 
otic fluid,  and  the  rhythmical  contractions  and  expansions  in 
the  amnion,  may  not  be  doubted  for  a  single  moment,  since  the 
former  would  increase  pressure,  while  the  latter  should  produce 
the  necessary  changes  in  pressure  in  the  embryo  for  compelling 
respiration  and  circulation  to  be  in  correspondence  with  the 
nutritive  and  functional  processes  in  the  growing  chick,  both 
of  which  are  constantly  extending  their  limits  and  requiring 
more  and  more  force  for  effecting  them.  These  slow  pulsations 
in  the  amnion  of  the  chick  answer  to  the  placental  and  uterine 
souffle  in  gestation,  the  principle  being  precisely  the  same. 
How  otherwise  explain  this  circumstance  %  But,  as  has  already 
been  remarked,  all  pulsations  relate  to  changes  in  pressure, 
and  these  pulsations  in  the  amnion,  together  with  the  amniotic 
fluid,  relate  to  changes  in  pressure  in  the  embryo  for  increasing 
circulation  of  the  juices. 

The  explanation  of  the  mechanics  is  sufficiently  easy  ;  nota- 
bly, there  are  two  cardinal  points  from  which  to  regard  it — one 
in  the  allantois,  the  other  in  the  embryo.  They  may  be  re- 
garded as  the  poles  to  this  circulation,  in  which  the  action 
alternates  for  assisting  the  circulation  by  rhythmical  changes 
in  pressure,  and  following  each  other  in  regular  order  and  suc- 
cession readily  produce  afflux  and  efflux  of  the  fluids  commen- 
surate with  the  physiological  requirements.  First,  commencing 
with  the  movement  of  expansion  in  the  allantois.  The  increase 
in  pressure  which  this  produces  in  the  embryo  through  the 
amniotic  fluid  occupying  more  room,  consequently  encroaching 
upon  the  amniotic  area,  should  cause  a  corresponding  amount 
of  the  venous  blood  to  flow  with  increased  energy  toward  the 
allantois,  the  point  of  low  pressure  within  the  egg  (the  heart 
and  vascular  system,  of  course,  assisting  in  this) ;  and  nice 
versa  during  contraction.  The  rhythmical  contractions  and 
expansions  in  the  amnion  have  the  effect  of  increasing  and 
diminishing  pressure  in  the  embryo  itself,  and  by  relieving 
pressure  in  the  allantois,   enables  this  to  expand  pari  passu 


346  PHYSIOLOGY    OF   THE   CIRCULATION". 

with,  contraction  in  the  amnion  for  aspirating  the  venous 
blood,  at  the  same  time  that  it  aspirates  the  air  through  the 
outer  membrane  and  pores  of  the  shell.  But  when  the  move- 
ment is  reversed  by  expansion  of  the  amnion,  the  reduction 
in  pressure  which  this  effects  in  the  embryo,  together  with 
the  simultaneous  increase  of  pressure  it  produces  in  the  allan- 
tois  by  forcibly  compressing  this  against  the  shell  wall,  causes 
the  aerated  blood  in  the  latter  to  flow  with  augmented  speed 
into  the  heart  of  the  embryo,  the  allantois,  itself,  also  partici- 
pating in  this  action,  the  fluid  it  contains  enabling  it  to  effect 
such  rhythmical  compression  of  the  capillary  plexuses  (Fig. 
14:5).  The  following  forcible  illustration  (Fig.  144)  by  the  dis- 
tinguished biologist  at  Jena  will  serve  to  impress  the  matter 
It  represents  early  stage  in  development  (third  week  in  gesta- 
tion) in  the  human  embryo,  it  will  be  seen  that  pressure  is 
increased  at  the  cardinal  points,  namely,  vitellus  (a),  the  body 
of  the  embryo  (c),  and  placenta  (5),  which  is  fundamentally 
the  same  as  the  allantois,  though  the  office  of  the  latter  is 
mainly  respiratory.  As  the  embryo  and  allantois  are  elabo- 
rated out  of  the  material  in  the  vitellus,  this  would  explain 
the  greater  accumulation  of  fluid  in  this  locality  for  com- 
pelling circulation  toward  those  two  points,  while  the  rhyth- 
mical contractions  of  the  yolk  sac  should  greatly  expedite  it. 
For  increasing  circulation  between  the  embryo  and  placenta 
(c,  b),  commensurate  pressure  is  produced  by  accumulation  of 
fluid  in  these  two  points  or  poles  of  this  circulation.  This, 
together  with  the  action  in  the  membranes  themselves,  and 
the  heart  and  vessels,  is  sufficient  for  carrying  on  circulation 
in  the  initial  stages  of  embryonic  evolution  ;  but  with  the  in- 
crease of  growth  comes  increasing  difficulty  for  effecting  it : 
hence  the  pumping  action  which  is  set  up  in  the  placenta  and 
womb,  together  with  the  accumulation  of  amniotic  fluid  for 
transmitting  these  actions  upon  the  embryo,  as  described 
above.  Thus,  everything  is  in  correspondence — the  liquor 
amnii,  the  increasing  growth  of  the  placenta  and  the  number 
of  muscles  and  nerves  in  the  walls  of  the  womb — and  so  con- 
tinues till  the  close  of  pregnancy.  In  other  words,  it  all  forms 
a  connected  whole  in  the  mechanics  of  the  embryonic  circu- 
lation.    The  absence  of  a  shell  wall  permits  expansion  in  the 


PHYSIOLOGY   OF  THE   CIRCULATION. 


347 


chorion  pari  passu  with  the  growth  of  the  embryo,  while  the 
womb  expands  in  concert  with  this  action  in  the  chorion  and 
embryo. 


Pig.  143.— Diagram  of  Young  Embryo  (Chick)  and  its  Vessels,  showing  circulation  of 
umbilical  vesicles,  and  also  that  of  allantois,  beginning  to  be  formed. — Dalton. 


Pig.  144. — Human  Embryo  in  the  Third  Week. — Haeckel.      «,  Large  globular  yolk  sac  ; 
b,  allantois  ;  c,  amnion  ;  d,  tufted  chorion.     There  are  yet  no  limbs. 

In  the  case  of  the  bird,  the  pumping  action  in  the  abdomen 
(the  soft  hinder  parts  of  the  bird)  for  pumping  air  and  blood 
through  the  alveoli,  is  set  up  in  the  latter  days  of  incubation, 
when  rapid  atrophic  changes  soon  separate  the  umbilical  ves- 
sels, and,  breaking  the  now  attenuated  and  fragile  shell  wall 


348 


PHYSIOLOGY   OF  TIIE   CIRCULATION. 


with  its  beak,  it  finally  makes  its  escape,  leaving  the  allantoic 
and  atrophied  membranes  behind. 


Fig.  145. — Ovum  from  the  Mule. — l,eydig.     u,  Nucleus  ;    b,  cell  body  ;    c,  thickened 
corpuscle  traversed  by  pores. 


Fig.  146. — The  Human  Egg  from  the  Ovary  of  the  Female  ;  much  enlarged. — Haeckel. 
The  entire  egg  is  a  simple,  globular  cell.  The  greater  part  of  the  spherical  egg-cell 
is  formed  by  the  egg-yolk,  or  the  granular  cell-substance  (protoplasm),  which  is  com- 
posed of  innumerable  delicate  yolk  granules,  with  a  little  intervening  substance.  The 
germ- vesicle,  answering  to  the  cell-kernel  (nucleus)  lies  in  the  upper  part  of  the  yolk. 
It  contains  a  dark  nucleolus  or  germ  spot.  The  globular  mass  of  yolk  is  suiTOunded  by 
a  thick  transparent  egg  membrane  {znna  jiellueida).  This  is  penetrated  by  the  pore- 
canals,  in  the  form  of  very  numerous  hair-like  Unes,  which  run  rapidly  toward  the 
centre  of  the  globe  ;  through  these  the  thread-shaped,  moving  sperm-cells  pass,  in 
the  process  of  impregnation,  into  the  egg-yolk. 

The  common  relation  which  animal  life  sustains  to  the  or- 
ganic laws  has  forcible  illustration  in  the  very  ova,  the  struc- 
ture being  fundamentally  the  same  (Figs  145,  146). 


CHAPTER  XV. 

ANIMAL  TEMPERATURE  AND  THE  NECESSITY  FOR  A  THERMIC 
CENTRE  IN  THE  MEDULLA  OBLONGATA. 

Body-Temperature — Why  the  Thermic  Centre  Should  be  Correlated  with  the  Respira- 
tory, Vaso-Motor  and  Voluntary-Motor  Centres  in  the  Medulla  Oblongata — Relations 
which  Respiration  Sustains  to  Metabolism — Ditto,  Metabolism  to  Body-  Temperature 
— The  Vaso-Motor  Centre  and  the  Vascular  Arrangements  in  the  Skin  for  Reducing 
Excessive  Body-Temperature  so  as  to  Maintain  a  Balance  in  Temperature  in  the 
Organism — Mode  of  Imitating  This  in  Febrile  Conditions  by  Therapeutical  Device  as 
the  Most  Effective  Means  of  Controlling  Body-Temperature,  Thereby  Conserving 
Life  and  Expediting  Recovery — Use  of  Respiratory  Sedatives  in  Connection  There- 
with, and  Rationale  of — Peculiarities  that  Obtain  in  Dogs  and  Birds  for  Facilitating 
Discharge  of  Body-Temperature  when  Excessive — Seat  of  Oxidation. 

Temperature  is  treated  more  advantageously  in  connection, 
with  development,  maintaining  mobility  among  the  molecules, 
for  which  a  special  physiological  adjustment  obtains  in  the 
organism,  but  which  differs  in  the  plants,  as  well  as  in  the 
animals,  though  the  oscillations  from  the  norm  in  any  one  of 
them  is  very  limited,  and  the  sliding  scale  in  the  warm- 
blooded may  be  roughly  estimated  at  from  six  to  ten  degrees 
only,  either  a  fall  or  rise  in  body-temperature  of  several 
degrees  speedily  bringing  life  to  an  end.  It  will  be  neces- 
sary, however,  to  briefly  refer  to  it  here  in  order  to  show  the 
relations  which  body-temperature  sustains  to  respiration  and 
circulation,  whereby  it  is  regulated  so  as  to  maintain  a  balance 
in  the  organism  ;  otherwise  impossible.  And  here  we  have  to 
mention  that  the  thermic  centre  for  the  organism  (which  is 
now  almost  universally  conceded  the  elaborate  researches  of 
Pfluger,  making  the  argument  unanswerable)  is  also  correlated 
with  the  other  nervous  centres  in  the  medulla  oblongata,  or 
respiratory,  vaso-motor,  voluntary-motor  and  thermic,  making 
four  in  all,  unless  the  trophic  be  also  included,  which  would 
seem  reasonable  and  natural,  since  nutrition  necessarily  de- 
pends upon  temperature  and  circulation. 


350   FOUR  NE1IVOUS  CENTRES  IN  THE  MEDULLA  OBLONGATA. 

The  reasons  upon  which  this  opinion  is  based  would  seem 
to  be  incontrovertible.  Briefly  summarized,  they  are  as  fol- 
lows : 

First  and  foremost,  the  scheme  calls  for  this  in  order  to  pro- 
duce continuity  in  force,  otherwise  impossible,  and  the  me- 
dulla oblongata  being  the  common  centre  of  nervous  force  for 
the  organism,  it  would  naturally  include  the  thermic  centre. 

2.  Since  heat  is  the  product  of  metabolism  or  the  nutritive 
and  chemical  changes  going  on  in  the  body,  but  principally 
the  latter,  Tjroduced  by  the  oxidizing  processes  generating 
carbonic  acid,  especially  in  the  muscles  during  musculation, 
and  the  commerce  is  pumped  through  the  tissues  for  the  pur- 
pose by  means  of  respiration  and  the  pumping  actions  in  the 
heart  and  vessels,  as  has  been  fully  shown  ;  it  follows  that  the 
thermic  centre  should  be  correlated  with  the  respiratory  and 
vaso  -motor  centres  for  compelling  prompt  response  in  order  to 
evolve  heat  in  the  measure  of  the  physiological  requirements, 
furnishing  the  requisite  fuel  for  the  purpose,  otherwise  im- 
possible ;  hence  the  correlation  of  the  thermic  with  these 
other  two  centres. 

3.  Active  musculation  is  the  principal  means  for  rapidly 
generating  heat  in  the  body ;  but  since  the  muscles  can  en- 
dure but  a  limited  amount  of  heat  above  the  norm,  which  is  93.5 
degrees  Fahr.  for  man,  it  follows  that  there  must  be  some  ready 
means  for  rapidly  bringing  the  blood  to  the  skin-surface  when 
body-temperature  is  excessive,  for  effecting  reduction  in  temper- 
ature by  means  of  radiation  and  evaporation  effected  by  means 
of  sweat,  which  is  poured  out  for  the  purpose,  the  vast  capil- 
lary network  in  the  skin  being  widely  expanded  for  the  pur- 
pose, producing  the  characteristic  flushing  of  the  skin  during 
active  exercise  ;  but  in  order  to  accomplish  these  results,  the 
thermic  centre  would  have  to  be  correlated  with  the  vaso- 
motor centre  for  compelling  prompt  response  in  the  vessels 
in  the  measure  of  the  requirements.  In  fine,  respiration, 
circulation  and  temperature  rise  and  fall  together,  rising  and 
falling  with  the  activities;  while  the  vaso-motor  centre,  by 
being  connected  with  the  thermic  centre,  compels  the  local 
actions  in  the  skin  for  maintaining  a  balance  in  tempera- 
ture ;  otherwise,  the  muscles  would  be  destroyed  in  their  own 


THE   AUTOMATIC   FUEISTACE   IN   THE   TISSUES.  351 

heat,  and  the  activities  would  be  speedily  fatal,  as  must  ap- 
pear obvious.  Hence,  the  correlation  of  the  thermic  centre 
with  the  respiratory,  vaso-motor  and  voluntary-motor  centres 
the  whole  being  woven  in  and  in,  and  must  be  so,  in  the  very 
nature  of  things.  Loss  in  body-temperature  is  announced  by 
sensory  impressions  in  the  skin ;  the  animal  shivers,  the 
teeth  chatter,  and  the  furnace  is  set  going  again  for  raising 
temperature,  the  animal  moving  about  restlessly  for  starting 
the  action  in  the  muscles,  doing  so  unconsciously,  hut  forced 
to  it  by  imperative  demand  in  the  organism.  This  hurries 
respiration  and  circulation  for  supplying  the  fuel,  which 
soon  puts  up  temperature  to  the  norm  again.  At  the  same 
time  the  appetite  is  increased,  while  the  digestive  and  assimi- 
lative processes  are  more  active,  and  from  these  combined 
sources  temperature  soon  rises  with  the  swell  in  the  activities, 
which  puts  up  respiration  and  circulation  correspondingly  ; 
so  that  prompt  help  comes  with  the  supply  of  food,  while 
force  and  fnel  are  economized,  if  the  animal  be  housed  so 
as  to  diminish  external  cold,  and  especially  if  protected  by 
suitable  clothing  for  diminishing  radiation  and  conduction, 
the  heavy  coat  of  hair  which  is  developed  with  the  accession 
of  cold  weather  subserving  this  function  in  the  animals,  while 
man  long  since  has  learned  the  art  to  perfection  by  raiding 
them  for  their  shaggy  coats,  at  the  same  time  aiding  himself 
with  fire  or  artificial  heat.  But  the  physiological  fact  to  be 
kept  uppermost  in  the  mind  is  the  correlation  of  the  great 
nervous  centres  in  the  medulla  oblongata,  making  respiration 
the  basis  of  all  the  movements,  while  the  exhalation  of  car- 
bonic acid  from  the  lungs  is,  so  to  speak,  the  smoke  of  the 
automatic  furnace,  the  principle  being  the  same  precisely  as 
obtains  in  a  grate  and  the  burning  of  coal;  only,  that  the 
tissues  subserve  the  purpose  of  a  grate,  the  fuel  furnished  by 
means  of  respiration  and  circulation,  while  the  pumping 
actions  invoke  the  law  of  pressure  which  applies  for  compel- 
ling supply  to  be  equal  to  demand.  All  plain  enough,  and 
easily  understood  from  the  stand-point  of  the  law  underlying 
the  organism  and  the  action  of  the  special  forces  which  apply 
in  the  case. 

The  increase  in  temperature  when  the  bodily  activities  are 


352  MAINTAINING    A    BALANCE   IN   TEMPERATURE. 

in  abeyance,  is  produced  by  prolonged  and  excessive  reflex  irri- 
tation propagated  from  the  mucous  surface  of  some  of  the  organs, 
or  from  the  skin,  or  by  blood-poison,  as  in  traumatism  and  the 
essential  fevers  acting  directly  upon  the  nervous  centres  of  res- 
piration and  circulation,  and  in  the  fact  of  retrograde  metamor- 
phosis from  loss  of  vitality  produced  by  the  disturbed  balance, 
the  supply  of  oxygen  brought  into  the  tissues  by  the  increased 
respiration  and  circulation  being  greatly  in  excess  of  the  de- 
mands, consequently  tending  to  excessive  oxidation  and  retro- 
grade metamorphosis,  whereby  the  tissues  are  consumed  and 
rapid  wasting  and  shrinkage  result  as  an  inevitable  sequence, 
while  the  matter  is  due  entirely  to  excessive  stimulation  of  the 
respiratory  and  vaso-motor  centres.  Take  the  temperature  and 
count  the  respirations  and  the  rhythms  in  the  heart  and  arteries, 
in  proof  of  this  statement.  Yes,  there  is  correspondence.  Now, 
then,  how  account  for  this  circumstance,  but  in  the  correla- 
lation  of  the  three  nervous  centres,  as  alleged  \  And  if  the 
respiratory  and  vaso-motor  are  in  the  medulla  oblongata,  it 
follows  that  the  thermic  centre  is  also  there,  for  in  no  other 
way  could  continuity  in  force  be  produced  for  effecting  this 
cycle  of  motion.  Then,  again,  when  temperature  falls, 
as  in  the  crisis,  respiration  and  circulation  are  in  corre- 
spondence, at  the  same  time  shifting  the  blood  to  the  surface 
in  order  to  cool  it ;  the  skin  is  reeking  with  perspiration  welling 
out  of  every  pore  for  carrying  off  temperature,  the  same  as  in 
bodily  exercise — an  heroic  effort  of  life  to  throw  off  the  toils 
in  which  she  is  caught  by  the  very  nature  of  her  framework, 
and  the  rude  experiences  to  which  the  animal  is  exposed.  It 
is  thought  by  some*  that  the  action  in  the  vaso-motor  centre 
would  explain  the  phenomena,  but  the  respiratory  should  be 
included  as  well,  so  as  to  diminish  the  supplies,  while  there 
must  be  a  heat-centre,  one  exceedingly  sensitive  to  heat,  for 
due  notification  of  local  increase  with  the  power  of  promptly 
compelling  the  requisite  vascular  changes  in  the  skin  and 
internal  parts  for  rapidly  expanding  the  external  and  con- 
tracting the  internal  vessels,  at  the  same  time  producing 
copious  perspiration ;  otherwise  life  would  suffer  speedy  ex- 

*  Pfluger's  Archiv.,  iii.  (1870,  504  ;  ibid.,  v.  (1872).  77.     Heidenhain. 


PHYSIOLOGY   OF    FEVFK.  853 

tinction  by  the  voluntary  movements  themselves,  not  to  men- 
tion the  reflex  irritations  and  the  action  of  blood-poison,  as 
must  appear  obvious.  It  follows  that  the  thermic  centre  must 
be  correlated  necessarily  with  the  other  centres  in  the  medulla 
oblongata,  the  solar  centre  for  the  organism.  Take  a  very 
simple,  as  also  a  very  common  case,  notably,  acute  tonsillitis 
in  the  child.  It  is  notorious  that  body-temperature  rushes 
up  with  extraordinary  rapidity  in  this  condition.  Why? 
The  explanation  is  easy.  Thus,  the  tonsils  are  in  close 
proximity  to  the  medulla  oblongata,  immediately  adjacent 
— you  might  say  right  in  front  of  it,  so  short  the  distance 
between  them ;  hence,  reflex  irritation  falls  at  once,  and 
with  all  of  its  force,  upon  the  respiratory  and  vaso-motor  cen- 
tres, and  the  increased  rhythms  which  this  produces  by  com- 
pelling excessive  importations  of  the  commerce  into  the 
organism  and  the  tissues  rush  up  the  temperature  corre- 
spondingly from  the  excessive  oxidation  it  induces.  In- 
deed, a  fretful  child  may  readily  induce  a  febrile  paroxysm 
at  any  time  by  a  violent  fit  of  crying,  but  the  sleep  which  fol- 
lows close  upon  exhaustion,  by  slowing  respiration  and  circu- 
lation, together  with  the  copious  diarjhoresis  which  accompa- 
nies it,  soon  reduces  temperature  to  the  norm  again,  so  that 
everything  runs  on  as  before.  The  practical  deduction  to  be 
drawn  therefrom  in  the  treatment  of  the  essential  fevers — to 
induce  sleep  artificially  by  means  of  chloral,  etc.,  at  the  same 
time  bathing  the  skin,  or  rather  keeping  the  surface  wet  by 
means  of  cloths  steeped  in  warm  water — e.  g.,  sponging  the 
night-clothes  with  this  so  as  to  obviate  shock  or  arouse  the 
patient ;  otherwise  inevitable.     More  of  this  presently. 

In  cases  of  extensive  burns  and  scalds,  which  are  always  fol- 
lowed by  increased  respiration  and  circulation,  producing 
fever,  how  explain  this  circumstance  if  not  by  reflex  irritation 
in  the  respiratory  and  vaso-motor  centres,  thereby  producing 
increased  respiration  and  circulation,  and  making  oxidation 
excessive  in  the  tissues  ?  The  very  principle  in  the  mechanism 
lays  the  whole  thing  bare.  The  office  of  therapeusis  is,  of 
course,  to  repress  the  irritation  by  appropriate  local  and  con- 
stitutional remedies,  or  by  soothing  applications  and  excluding 
the  air,  at  the  same  time  administering  the  salts  of  morphia, 


354  PHYSIOLOGY   OF   FEVER. 

etc.,  supporting  the  patient.  In  the  case  of  blood-poisoning, 
permanent  relief  comes  only  by  eliminating  the  materies  morbi 
through  the  special  emunctories  ;  Medicine,  as  it  were,  sitting 
by  the  tiller  and  guiding  the  life-boat  so  as  to  keep  it  off  the 
rocks  and  dangerous  shoals,  clear  of  the  breakers  and  in  the 
deep  water,  by  keeping  down  excessive  temperature  by  respira- 
tory sedatives  for  reducing  the  rhythms  in  the  lungs  and 
circulatory  apparatus,  at  the  same  time  reducing  temperature 
in  the  manner  as  stated,  repeatedly  applying  the  thermometer 
to  learn  the  state  of  the  temperature,  saving  the  patient  by 
constant  vigilance.  And  Medicine  is  made  a  helpmate,  not  an 
executioner,  for  struggling  life.  The  intimate  relations  which 
the  skin  sustains  to  the  respiratory  and  vaso-motor  centres,  by 
reason  of  the  sensory  nerves,  would  account  for  fever  induced 
by  irritations  of  any  kind,  e.  g.,  the  exanthemata,  which 
independent  of  the  special  action  of  the  materies  morbi  upon 
the  nervous  centres,  interfere  with  transpiration,  consequently 
producing  an  accumulation  of  heat,  as  it  were,  damming  it  in 
the  deap  vessels,  thereby  preventing  its  due  escape  from  the 
skin-surface.  The  high  temperature  in  this  class  of  cases  is 
undoubtedly  due  to  accelerated  respiration  from  excessive 
stimulation  of  the  respiratory  centre  from  reflex  actions  in  the 
skin,  together  with  the  suppression  of  the  secretory  functions 
in  the  skin  for  carrying  off  excessive  temperature.  The  role 
in  therapeusis,  however,  is  clear — namely,  repress  excessive 
temperature  in  the  manner  as  stated.  Small-pox,  so  malig- 
nant in  times  past,  when  the  patients  were  treated  in  close 
rooms,  but  now  in  open  wards  and  tents,  where  freest  ingress 
of  air  is  insured,  reduces  body-temperature  by  conduction, 
with  very  little  mortality.  In  all  these  cases  there  is  rapid 
respiration,  induced  by  excessive  irritation  in  the  skin,  com- 
bined with  the  direct  action  of  the  materies  morbi  upon  the 
respiratory  and  vaso-motor  centres.  In  addition  to  frequent 
bathing  of  the  skin,  for  supplementing  perspiration,  some 
respiratory  sedative,  so  as  to  induce  sleep,  e.  g.,  quinine, 
chloral,  etc.,  should  be  administered  in  cases  of  alarming 
temperature,  as  where  it  exceeds  105°  Fahr.  In  sleep,  tem- 
perature falls  because  respiration  falls  in  frequency.  And 
convalescence  in  fever  is  ushered  in  by  sleep;    sleep  and 


PHYSIOLOGY   OF    FEVEK.  355 

diaphoresis,  the  slowing  of  respiration  and  the  escape  of  heat 
by  evaporation  putting  an  end  to  the  fever. 

The  explanation  of  the  fall  in  temperature  produced  by  the 
slowing  of  the  respiratory  rhythms  is  obvious  enough.  Thus, 
taking  the  average  in  man,  sixteen  respiratory  rhythms  are 
sufficient  for  maintaining  the  body-temperature  at  98. 5  degrees 
Fahr. ,  each  volume  of  air  inspired  representing,  of  course,  a 
given  volume  of  oxygen,  which  is  sufficient  for  maintain- 
ing the  norm.  In  other  words,  sixteen  pumps  of  the  respi- 
ratory mechanism  (which,  of  course,  would  include  the  ac- 
tions in  the  heart  and  blood-vessels,  since  it  all  forms  a  con- 
nected movement)  are  sufficient  for  maintaining  the  body-tem- 
perature at  98.5  degrees  Fahr.  Now,  then,  any  circumstance 
which  should  increase  the  rhythms  would  produce  corre- 
sponding elevation  of  the  temperature,  but  for  the  compre- 
hensive arrangements  that  obtain  for  discharging  heat  from 
the  body-surface  for  maintaining  a  balance,  already  referred 
to  ;  but  in  abnormal  conditions  this  is  more  or  less  inter- 
rupted. 

And  for  that  reason  the  public  speaker  "warms  up"  as  he 
proceeds  in  his  subject,  speaking  more  and  more  rapidly,  tem- 
perature rising  correspondingly. 

It  is  the  same  in  the  vocalist,  as  i  have  proven  over  and  over 
again,  by  taking  the  temperature  before  the  commencement 
of  the  exercise,  during  it,  and  afterward,  and  while  the  tem- 
perature varied  somewhat,  always  finding  it  higher  than  be- 
fore, from  one  to  two  degrees,  according  to  the  energy  in  the 
respiratory  rhythms.  When  the  temperature  increases  from 
one  to  two  degrees,  the  balance  is  struck  by  reason  of  the  action 
in  the  skin  ;  else  he  must  stop,  from  exhaustion  induced  by  the 
action  of  temperature  upon  the  nervous  apparatus,  which  is 
Nature' s  method  for  putting  an  end  to  the  exercise.  But  muscu- 
lar exertion  of  any  kind  produces  increase  in  temperature  !  Of 
course  it  does  ;  but  is  not  respiration  in  correspondence?  So, 
likewise,  the  action  induced  by  cold,  since  this  rapidly  with- 
draws heat,  and  respiration  must  be  hurried  correspondingly 
for  maintaining  a  balance  ;  otherwise  impossible.  And  when 
the  air  is  loaded  with  moisture  which  rapidly  absorbs  body- 
temperature,  the  individual  exposed  to  it  had  better  keep  the 


366  NECESSITY    FOR   A   THEItMIC    CENTRE. 

muscles  in  action  for  rapidly  evolving  heat,  and  he  "sits"  at 
his  peril.  For  dismissing  heat,  the  vessels  in  the  skin  are  ex- 
panded, and  the  deep  vessels  contracted ;  but  for  retain- 
ing it,  the  surface  vessels  are  contracted,  and  those  in  the 
deep  territories  are  expanded  correspondingly,  in  order  to 
shift  the  blood  from  one  to  the  other  part,  at  the  same  time 
increasing  respiration  for  supplying  the  additional  quantity 
of  fuel  and  oxygen  which  are  called  for  for  evolving  heat  and 
maintaining  a  balance  in  the  organism,  otherwise  impossible. 

Thus,  we  have  not  only  the  vaso-motor,  but  the  respiratory 
centre  included  in  the  thermal  mechanics  for  increasing  oxida- 
tion. When  the  loss  in  heat  is  rapid,  as  when  cold  is  ex- 
cessive, the  appetite,  digestive  and  assimilative  processes  are 
increased,  as  has  already  been  remarked — the  latter  from 
the  additional  amount  of  blood  which  is  sent  to  the  organs 
from  the  external  parts — so  that  the  whole  mechanics  for 
producing  temperature  work  together  in  perfect  concord  and 
unity.  But  for  its  accomplishment,  there  must  be  a  ther- 
mic centre  for  receiving  and  transmitting  the  sensory  im- 
pressions produced  by  heat  and  cold,  and  for  compelling  the 
requisite  vascular  and  respiratory  movements  suitable  to 
the  occasion.  In  other  words,  a  special  sense  necessitates  <ni 
appropriate  nervous  reflex  centre  for  duly  impressing  the  ap- 
pellate nervous  centres  for  compelling  the  requisite  move- 
ments in  the  mechanics  for  maintaining  a  balance  in  the  organ- 
ism, otherwise  impossible.  And  when  one  reflects  upon  the 
preeminent  importance  of  temperature  in  the  animal  organism, 
the  occasion  for  this  comprehensive  arrangement  for  produc- 
ing and  regulating  it  will  at  once  appear  obvious. 

Finally,  the  experiments  of  Naunyn  and  Quincke*  show 
that  whether  division  be  made  above  or  below  the  medulla 
oblongata,  it  is  attended  by  a  rise  in  temperature,  proving 
that  the  thermic  centre  is  in  the  medulla  oblongata,  while  the 
very  argument  used  by  Schroflf  to  prove  the  circumstance 
was  due  to  fever  occasioned  by  the  mere  wound  itself,  but 
confirms  it,  since  it  shows  the  nervous  centres  in  the  medulla 
oblongata  can  produce  fever,  and  since  fever  is  the  result  of 

*  DuBois-Raymond's  Archiv.,  1866.  p.  151  ;  1869.  pp.  174,  521. 
f  Wien.   Sitzungsherichte,  Lx.  xiii.  (1876). 


THERAPEUSIS    OF   FEVER.  357 

excessive  metabolism,  it  would  show  that  by  simply  stimulat- 
ing the  respiratory  would  necessarily  involve  the  vaso-motor 
and  thermic  centres. 

So,  then,  we  can  readily  understand  why  respiration  should 
be  in  correspondence  with  metabolism,  both  in  normal  and 
abnormal  conditions  of  the  system,  and.  why  it  should  be  more 
rapid  in  warm  than  in  cold-blooded  animals.  Finally,  why  it 
should  oscillate  with  external  temperature,  and  with  the 
states  of  activity  and  repose. 

An  Important  Medical  Case,  Showing  the  most  Effective  and 
Kindly  Means  for  Rapidly  Reducing  Excessive  Body- 
Tem/perature. 

The  following  deeply  interesting  medical  case,  which  came 
within  my  own  knowledge,  may  prove  of  great  practical 
use  to  the  profession,  and  since  this  is  the  chief  object 
sought  to  be  attained  in  physiological  inquiry,  it  should 
entitle  it  to  a  place  in  the  text.  It  occurred  in  connec- 
tion with  acute  inflammatory  rheumatism,  with  so-called 
metastasis  to  the  brain,  and  known  as  "cerebral  rheu 
matism"  ;  but,  in  point  of  fact,  is  the  result  simply  of 
excessive  blood-temperature.  It  was  produced  in  a  lad  thir- 
teen years  of  age  (son  of  a  prominent  jurist),  from  exces- 
sive bathing,  followed  by  high  fever  and  pain  in  the  joints  for 
the  first  several  days,  then  the  brain-symptoms  spoken  of  set 
in.  When  called  in  consultation,  I  found  the  patient  per- 
fectly unconscious  ;  comatose ;  the  extremities  cold,  but  the 
skin  over  trunk  above  the  normal  temperature  ;  in  the  rectum, 
10&2  Fahr.  ;  pulse  168;  respirations,  34  per  minute.  The 
cerebral  symptoms  had  set  in  sometime  during  the  night  and 
early  morning,  ushered  in  by  delirium.  The  treatment  recom- 
mended above  was  adopted.  All  the  covering  save  his  night- 
shirt was  removed,  and  this  was  now  sponged  with  lukewarm 
water,  while  for  facilitating  evaporation  fanning  was  resorted 
to.  It  is  astonishing  how  rapidly  the  body- temperature  is 
reduced  in  this  manner.  In  something  like  two  and  one-half 
hours  the  temperature  had  fallen  to  103  degrees  Fahr.,  with 
marked  improvement  in  all  the  symptoms  ;  pulse  130 ;  respi- 
rations. 28  per  minute,  with  partial  return  to  consciousness, 


358  thek.apkusis  of  fever. 

showing  the  comatose  condition  had  been  superinduced  by 
excessive  blood-temperature  The  sponging  of  the  skin,  or 
rather  the  shirt,  was  now  discontinued,  but  he  took  five  grains 
of  sulphate  of  quinine  very  readily,  and  swallowed  it  down 
with  water,  his  hand  to  the  glass  regulating  it.  We  then  left 
him  for  several  hours,  but  leaving  strict  orders  to  watch  the 
temperature,  else  he  would  have  a  relapse,  the  head-symp- 
toms would  return.  During  our  absence,  however,  complain- 
ing of  cold,  his  mother  wrapped  him  in  blankets,  his  shirt 
also  was  changed,  and  when  we  returned,  to  my  great  con- 
sternation and  alarm,  he  was  again  unconscious  and  breathing 
rapidly  ;  the  pulse  accelerated ;  while  the  thermometer  in  the 
rectum  registered  107  degrees  Fahr.,  or  nearly  as  high  as  at  first. 
The  treatment  by  sponging  was,  of  course,  at  once  resumed, 
and,  fortunately,  in  the  course  of  three  hours  the  temperature 
fell  to  10:5.1  degrees  Fahr.,  with  great  improvement  in  the 
symptoms  ;  respiration  27,  pulse  138  per  minute,  with  return 
to  consciousness,  and  proving  conclusively  that  the  coma 
had  been  induced  by  the  excessive  blood- temperature.  .After 
this  he  was  carefully  watched,  and  made  a  good  recovery  in 
the  course  of  four  or  five  weeks,  the  articular  affection,  which 
returned  almost  at  once,  running  its  usual  course. 

By  thus  throwing  wide  open  the  escape-valve,  so  to  speak, 
to  body-temperature  by  the  free  use  of  water  to  the  skin,  to- 
gether with  the  use  of  respiratory  sedatives  to  inhibit  excess- 
ive importations  of  oxygen,  constitutes  the  most  efficient 
means  for  controlling  body-temperature.  In  respect  to  the 
free  use  of  whisky,  which  is  very  beneficial,  it  acts  in  two 
directions  :  First,  by  springing  open  the  skin  capillaries  it  de- 
termines the  blood  to  the  surface,  thereby  cooling  it.  It  also  acts 
as  a  respiratory  sedative  and  tends  to  stay  metabolism.  By 
producing  expansion  in  the  capillary  network  of  the  skin  it 
tends  to  promote  diaphoresis,  which  accelerates  the  discharge 
of  heat.  There  is  nothing  better,  therefore,  than  this  supple- 
mental treatment  for  controlling  temperature  in  low  febrile 
conditions.  I  think  the  benefit  derived  from  quinine  has  simi- 
lar explanation,  expanding  the  surface  capillaries  and  inducing 
sleep,  but  large  doses  are  required  to  make  it  effective.  in 
traumatism  the  preparations  of  opium,  by  reducing  sensibility 


THEKAPEUSIS    OF   FEVER.  '659 

and  tims  inhibiting  reflex  action,  come  in  well.    It  is  needless 
to  extend  the  matter. 

One  thing  more,  however,  before  we  bring  this  matter  to  a 
close  :  The  Germans  (to  whom  we  are  indebted  for  the  "  cold- 
water  treatment"  in  fever)  are  undoubtedly  correct  in  principle, 
but  the  method  adopted,  or  that  by  the  cold  douche,  may  be 
carried  out  successfully  in  hospitals,  but  it  will  not  answer  in 
private  practice  for  the  following  reasons  :  1.  The  prejudices 
of  the  patient  and  his  friends  are  against  it,  and  would  have  to 
be  borne  down  by  the  strenuous  efforts  of  the  physician,  and 
should  the  case  issue  fatally,  adverse  criticism  would  be  rife  ; 
not  to  mention  the  labor  it  involves  and  the  difficulty  of  ob- 
taining the  requisite  nurses  and  appliances.  2.  Infants  and 
young  patients  are  nearly  thrown  into  spasm  with  terror,  the 
shock  being  so  violent.  Whereas,  by  the  more  kindly  method, 
prejudice  will  not  seek  to  balk  you,  in  place  will  eagerly  assist 
you,  while  the  patient  is  soothed  and  his  high  temperature 
filched,  so  to  speak,  from  him  without  his  knowledge. 

n  animals  in  which  transpiration  is  not  so  abundant  as  in 
man — dogs,  for  example — compensation  is  made  through 
evaporation  from  the  mouth  aud  bronchial  mucous  mem- 
brane, when  temperature  is  excessive.  Notably,  the  animal 
throws  open  and  projects  the  tongue  far  out  of  the  mouth 
and  breathes  very  rapidly,  but  very  shallow,  so  as  to  obviate 
penetration  of  the  air  to  the  alveoli  as  much  as  possible, 
which  surges  in  and  out  of  the  mouth  and  proximal  por- 
tions of  the  trachea  and  bronchi,  and  volume  after  vol- 
ume of  aqueous  vapor  is  thus  discharged,  while  the  tongue 
fairly  smokes  with  the  fog  arising  from  it.  But  he  plunges 
into  water  the  first  opportunity  that  presents,  for  expediting  the 
process.  Birds  have  a  similar  action  in  the  throat,  which 
pumps  as  in  the  frog,  at  the  same  time  spreading  their 
wings,  so  as  to  facilitate  conduction  and  radiation,  seeking  the 
cool  places  and  burying  themselves  in  the  earth  when  they 
can,  scratching  holes  for  the  purpose. 

Concerning  the  Locality  where  Oxidation  is  Effected  — It  is 
believed  in  certain  quarters  that  oxidation  of  the  carbon  com- 
pounds by  means  of  which  force  is  evolved  and  carbonic  acid  is 
formed,  takes  place  principally  within  the  capillaries  ;  but  this 


360  SEAT  OF   OXIDATION. 

cannot  be,  for  the  reason  that  the  cell-brood  is  the  objective 
point  for  all  the  commerce,  as  they  are  the  workmen  in  the  tis- 
sues, therefore  the  seat  of  metabolism,  and  to  stop  short  of  its 
destination  would  defeat  the  scheme  in  the  circulation,  the  ves- 
sels being  a  carrier  simply,  but  under  control  of  the  cell-brood. 
In  other  words,  they  live  upon  the  stream,  and  withdraw  the 
commerce  as  it  is  needed  in  the  functions  and  for  repair  of  the 
tissues.  Furthermore,  the  fluids  flowing  from  the  tissue-inter- 
stices into  the  capillaries  (through  the  stomata  in  the  latter  dur- 
ing expansion),  are  waste  'products  that  are  pushed  at  once  into 
the  venous  system  by  the  subsequent  contraction  in  the  vessels 
and  the  pressure  in  the  arterial  system,  hence  have  no  oppor- 
tunity to  mix  with  the  arterial  blood  for  effecting  oxidation. 
Then,  again,  if  the  blood  itself  were  a  source  of  heat,  the  fact 
should  be  announced  in  the  lungs,  where  oxygen  mixes  freely 
with  the  venous  blood ;  nevertheless,  it  has  been  proven  to 
demonstration  that  the  blood  in  the  left  side  of  the  heart  has  a 
lower  temperature  than  the  right,  which  is  heated  by  the  por- 
tal blood,  and  suffers  actual  loss  in  the  alveoli,  by  reason  of  con- 
duction and  an  amount  of  evaporation  from  the  mucous  surface. 
In  fine,  from  the  very  nature  of  the  mechanics,  the  cell-brood 
must  be  the  seat  of  metabolism,  respiring  through  the  vessels 
which  serve  to  connect  them  with  the  lungs  and  the  environ- 
ment, while  respiration  itself  is  increased  or  diminished  in 
correspondence  with  their  requirements.  It  is  very  compre- 
hensive, but  easily  understood. 


CHAPTER    XVI. 

THE  GREAT  ROLE  OF  CARBONIC  ACID  IN  THE  ANIMAL  ORGAN- 
ISM— A  FOOD  FOR  THE  TISSUES  AND  A  STIMULUS  FOK  THE 
FUNCTIONS. 

Functions  of  Carbonic  Acid  and  Nitrogen  Gases  in  Arterial  Blood — Explanation  for  the 
Disappearance  of  Carbonic  Acid  and  Nitrogen  in  Respiration,  the  Amount  of 
Carbonic  Acid  Expired  not  Representing  the  Chemical  Equivalent  of  the  Oxygen 
Inspired,  while  Nitrogen  in  Small  Amount  also  Disappears — Carbonic  Acid  and 
Nitrogen  Gases  Normal  Constituents  of  Arterial  Blood — Connection  with  the  Func- 
tions in  the  Intestines — Ditto  the  Nutritive  Processes — Its  Universal  Diffusion 
in  the  Air,  "Water,  Floral  and  Animal  Juices  and  Tissues — The  Universal  Appetite 
for  Carbonic  Acid  Drinks — A  Stimulus  to  the  Digestive,  Respiratory,  and  Vaso- 
Motor  Centres — Significance  of  Sleep — The  Odor  in  Human  Faeces — Genesis  and 
Functions  of  the  Gases  in  the  Air-Bladders  in  the  Fishes. 

The  fact  that  an  amount  of  carbonic  acid  and  nitrogen  dis- 
appears in  respiration,  that  the  quantity  of  carbonic  acid  ex- 
pired does  not  represent  the  chemical  equivalent  of  the  oxygen 
inspired,  the  same  remark  applying  to  nitrogen,  is  well  known 
and  commented  upon,  but  no  practical  inference,  or  one  re- 
lating to  special  functions  subserved  by  them,  drawn  from  it, 
but  regarded  as  a  curious  circumstance  only.  But  the  very 
fact  that  they  are  normal  constituents  of  arterial  blcod,'\ 
therefore  freely  distributed  through  the  organism,  is  of  itself 
sufficient  to  prove  they  must  subserve  important  uses  in  the 
functions,  and  are  not  to  be  regarded  as  interlopers,  so  to  speak, 
and  footpads  bent  on  mischief  and  all  possible  harm.  The 
only  deduction  made  being  that  the  quantity  of  gases  in 
the  blood  and  tissue-juices  is  regulated  by  what  is  known 
as  ' '  tension' '  in  the  gases  ;  but  what  this  tension  itself  refers 
to  is  not  stated,  and  we  are  left  as  much  in  the  dark  in 
the  one  as  in  the  other.  But  as  everything  in  the  body  has 
its  uses — moreover,  is  regulated  by  organic  laws,  which  are 
put  in  force  in  the  measure  of  the  physiological  requirements 
in  the  organs  and  tissues — we  may  rest  assured  that  philosophic 


3G2     CAKEONIC  ACID  and  nitrogen  gases  in  the  blood. 

scientific  reasons  may  be  given  for  their  presence  also,  with 
every  other  definite  arrangement  that  obtains,  and  it  cannot 
be  doubted  for  a  single  moment  even.  Moreover,  it  is  sus- 
ceptible of  easy  explanation,  as  we  shall  now  proceed  to  show, 
notably  : 

1.  From  the  very  nature  of  the  mechanics  in  the  intes- 
tinal canal,  it  is  easy  to  perceive  why  the  blood  is  charged 
with  the  gases,  since  this  is  necessary  for  increasing  pressure 
in  the  intestines  in  connection  with  the  digestive  and  absorp- 
tive functions,  and  for  maintaining  a  balance  in  the  portal 
circulation,  otherwise  impossible,  the  intestinal  gases  being 
increased  with  the  difficulties  in  this  circulation,  notably  in 
corpulency  (pp.  187-100) ;  and  since  pressure  is  constantly 
changing  in  correspondence  with  the  exigencies  in  the  func- 
tions, it  follows  that  some  comprehensive  arrangement  must 
obtain  for  secreting  and  absorbing  the  gases  in  the  organs  in 
the  measure  of  the  physiological  requirements  ;  hence  the  cir- 
cumstance that  carbonic  acid  and  nitrogen  gases  are  normal 
constituents  of  arterial  blood,  together  with  the  special  arrange- 
ments that  obtain  for  circulating  them  in  the  blood. 

During  digestion,  however,  the  amount  of  the  gases  expired 
is  in  excess  of  the  gases  inspired,  which  is  due  to  the  fact 
that  supply  is  greater  than  demand,  the  quantity  ingested  with 
the  food  and  others  generated  in  the  chemical  reactions  during 
digestion  being  more  than  sufficient  for  producing  only  a  given 
amount  of  pressure  in  the  intestines,  in  connection  with  the 
digestive  and  absorptive  processes,  otherwise  impossible ; 
since  the  gases  already  in  the  organs  when  food  is  ingested, 
together  with  that  carried  in  with  the  aliment,  and  generated 
by  the  chemical  reactions,  must  inevitably  produce  dangerous 
accumulations,  but  for  this  absorption  by  the  blood-vessels 
and  discharge  through  the  lungs ;  moreover,  if  this  were  not 
as  it  is.  it  would  defeat  the  purpose  of  the  arrangement  by  pre- 
venting the  due  amount  of  ingesta  ;  also  producing  pain  and 
discomfort,  besides  inducing  abnormal  changes.  Hence  this 
absorption  of  the  gases  and  their  discharge  through  the  lungs, 
the  most  expeditious  way  of  getting  rid  of  them. 

In  this  manner,  then,  pressure  in  the  intestines  is  easily 
regulated.    And  it  must  be  borne  in  mind  that  there  is  a  norm 


CAKBONIC  ACID  AND  JSTI'l  ROGEN  GASES  IN  THE  BLOOD.      363 

in  pressure  in  the  intestines,  and  that  the  object  of  peristalsis 
is  to  increase  or  diminish  it  as  occasion  may  require  in  the 
exigencies  in  the  functions  simply,  by  expanding  and  con- 
tracting upon  it  in  the  manner  as  stated  when  treating  of  the 
functions  in  the  intestines,  and  the  mechanical  action  of  the 
air-cushion  in  connection  therewith  (pp.  1^7-226).  This,  then, 
would  explain  that  circumstance,  and  it  can  be  explained  in  no 
other  way. 

2.  In  the  second  place,  carbonic  acid  functions  as  a  stimulus 
to  the  digestive  and  nutritive  processes,  promoting  rapid  di- 
gestion and  nutrition,  the  importance  of  which  it  would  be  diffi- 
cult to  overestimate.  They  can  be  treated  more  advantageously 
under  separate  heads. 

The  following  table  by  Bert  will  show  the  amount  and  rela- 
tive proportion  of  the  gases  in  arterial  blood : 

Carbonic  acid,  Carbonic  acid,  Total  gas 

disengaged  in  combi-  Carbonic  acid,  in  volume 

Oxygen,   by  a  "vacuum.  nation.                   total.  Nitrogen.       per  100. 

Arterial  blood.  15 -03           2799  1-15                   29"14  1-60  45-77 

Venous  blood..   8"17           31-27  2-38                   33-65  1-37  43-19 

It  will  be  seen  from  the  above  table  that  the  venous  blood, 
in  its  passage  through  the  lungs,  yields  up  less  than  one- 
fourth  of  the  free  carbonic  acid  contained  in  it,  the  greater 
portion,  or  more  than  three-fourths,  passing  on  into  the  arte- 
rial system.  Now,  then,  as  this  is  the  rule,  the  normal  con- 
dition, it  follows  that  carbonic  acid  gas  is  essential  to  the 
animal  organism.  A  bald,  naked  fact  of  enormous  import 
stands  out  there.  What  will  you  do  with  it  %  Shall  we  write 
underneath:  "Carbonic  acid  is  a  poison"?  You  may,  but 
I  never  shall ;  and  I  enter  my  solemn  protest  against  your 
doing  so. 

The  universal  appetite  for  carbonic  acid  drinks,  and  why  all 
natural  waters  are  saturated  with  id,  must  also  be  explained 
along  with  the  rest,  for  there  is  interdependence  and  connec- 
tion in  the  phenomena.  And  the  only  poison  I  see  is  in  the 
philosophy,  for  never  was  great  virtue  so  traduced  and  vili- 
fied, maligned  and  made  hideous  beyond  all  recognition  as  in 
this  case.  Because  carbonic  acid  does  not  support  combus- 
tion and  animal  life  cannot  therefore  live  in  it,  going  out  like 
a  candle  by  reason  of  the  absence  of  oxygen,  then,  forsooth,  it 


364  CA  HBO  NIC    ACID   NOT   A   POISON. 

is  placarded  "  a  deadly  poison."  A  "  negative  poison,''  then? 
No  !  iN  o  poison  whatever,  any  more  than  water  is  a  poison,  and 
for  the  same  reason  that  it  does  not  support  combustion.  " A 
coroners  jury"  may  have  such  opinion,  but  it  is  not  admissi- 
ble in  science.  Submerging  an  air-breather  in  carbonic  acid 
gas  speedily  puts  an  end  to  life,  but  not  more  speedily  than 
water,  both  producing  death  by  asphyxia  simply  ;  while 
simply  excluding  the  air  by  means  of  a  "gag"  would  have  the 
same  effect — producing  death  by  asphyxia.  But  would  you 
call  the  "  gag  "  a  poison  by  reason  of  that  fact  ?  "A  coroner's 
jury"  might,  forming  judgment  from  surface  appearances 
simply,  as  in  the  case  of  the  still  more  widely  circulated  but 
exploded  opinion  that  "  The  sun  rises  in  the  east  and  sets  in 
the  west."  But  to  put  such  stuff  in  scientific  works  is  enough 
to  run  one  mad.  (Ah  me  !  Mind  and  heart  alike  are  sveary.) 
This,  too,  when  arterial  blood  is  nearly  one-half  carbonic  acid 
gas  in  volume,  or  44.14  in  the  100 — more  in  reality  than  in 
venous  blood,  which  is  41.82  in  the  100 — while  all  the  juices 
and  tissues  are  literally  saturated  with  it,  and  all  natural 
waters  as  well ;  so  that  there  is  obviously  a  preconcerted  effort 
on  the  part  of  Nature  not  only  to  shut  off  all  escape,  but  to 
keep  the  tissues  saturated  with  it.  And  man,  instinctively 
cooperating  all  the  while,  demanding  carbonic  acid  in  unusual 
quantities  in  his  beverages— notably,  soda  water,  beer,  cham- 
pagne etc. — and  should  it  prematurely  escape  from  any  one 
of  them  from  untimely  uncorking,  making  it  "flat"  and  "un- 
palatable," is  spewed  out  of  the  mouth  as  worthless  ;  while 
water  which  has  been  boiled,  driving  the  gas  out  of  it,  is  a 
nauseous  dose.  And  everything  rushes  to  the  fountain  where 
the  waters  issue  from  the  earth,  and  most  highly  charged  with 
carbonic  acid,  giving  them  their  sparkling  appearance  and 
making  them  more  palatable,  at  the  same  time  also  serving  as 
the  most  effective  solvent  for  the  alkaline  earths  and  minerals. 
And  if  carbonic  acid  gas  be  a  poison,  as  alleged,  why  all  this  ? 
and  if  but  a  waste  product  in  arterial  blood,  in  the  name  of 
the  All !  why  are  there  not  more  emunctories  for  maintaining 
cleanliness  and  purifying  the  blood  of  carbonic  acid,  the  only 
exception  being  in  this  ?  What  a  mess  they  would  make  of  it ! 
Finally,  the  delightful  sensations  the  gas  produces   in  the 


FUNCTIONS  OF  CARBONIC  ACID  IN  ANIMAL  ORGANISMS.    365 

gullet  and  stomach  when  swallowed  shows  it  to  be  a  kindly 
and  effective  stimulant  to  the  digestive  processes. 

So,  then,  it  will  readily  be  perceived  that  to  dub  carbonic 
acid  gas  a  poison,  is  illogical  and  unscientific. 

And  when  men  seek  low  places  in  the  earth,  let  them  see  to 
it  that  carbonic  acid  is  not  in  there,  displacing  the  atmosphere^ 
by  its  greater  weight  and  ready  to  destroy  them  by  asphyxia. 

From  what  has  preceded,  it  will  be  readily  inferred  that  the 
carbonic  acid  secreted  in  the  stomach  and  intestines  during 
the  digestive  processes  subserves  not  only  the  mechanical 
uses  for  producing  the  changes  of  pressure  in  the  gut,  but 
at  the  same  time  it  acts  as  a  stimulus  to  the  secretory  and 
absorptive  processes  as  well ;  thereby  giving  another  manifes- 
tation of  the  comprehensiveness  in  animal  mechanics .  which  may 
be  seen  from  every  aspect  to  the  open  eye  and  thoughtful  mind. 

There  is  nothing  narrow  or  contracted  in  Nature,  but  broad, 
generous  and  wonderfully  comprehensive,  reaching  out  into 
unfathomable  space  and  distance  immeasurable,  while  method 
and  order  everywhere  obtain  ;  so  that  we  need  not  fear  to  ex- 
ceed her  resources  in  following  to  their  logical  results  the 
action  of  her  laws  in  living  organisms,  knowing  full  well  that 
everything  in  the  visible  universe  is  based  upon  law. 

Concerning  the  Relations  Which  Carbonic  Acid  Sustains 
to  the  Nutritive  Processes  in  Animal  Organisms. — It  is 
notorious  that  nutrition  is  more  active  during  sleep,  or  the 
period  when  carbonic  acid  tends  to  accumulate  in  the  blood 
and  tissues  from  diminished  respiration,  at  the  same  time 
there  is  tendency  to  venous  stasis  in  the  systemic  capillaries 
from  venous  obstruction  at  the  right  side  of  the  heart,  so  that 
when  very  profound  in  obese  individuals  there  is  a  degree  of 
lividity  in  the  skin,  especially  in  the  face  and  extremities :; 
and  when  produced  by  soporifics,  and  respiration  is  still  more 
slowed,  the  whole  surface  is  suffused  with  venous  discolora- 
tions.  Nevertheless,  during  this  condition  of  venosity,  nutri- 
tion is  very  active.  Why  ?  Undoubtedly  a  principle  is  in- 
volved, for  there  is  method  in  it.  Now,  then,  the  question: 
What  is  this  principle  which  applies  to  the  nutritive  processes  % 
We  know  the  one  which  applies  for  evolving  force  in  the 
organism,  and  it  remains  to  discover  the  one  which  applies  to> 


366  CARBONIC   ACID   A   FOOD   FOR  THE    TISSUES. 

nutrition ;  the  one  tending  to  dispersion,  the  other  to  accre- 
tion, or  growth ;  therefore  representing  opposite  conditions. 
The  circumstance  which  gives  the  clue  here  is  the  fact  that 
the  tissues  are  composed  of  compounds  of  carbon,  floral  as  well 
as  the  animal  tissues.  Now,  then,  put  these  two  facts  to- 
gether, placing  them  side  by  side,  for  they  are  complementary 
and  belong  together,  animal  resting  upon  floral  life,  morever, 
is  composite,  floral  structure  and  the  principle  in  floral  life 
pervading  it,  namely  :  1.  Carbonic  acid  is  the  principal  source 
of  floral  structure.  2.  The  most  active  nutrition  in  animal  life 
is  coincident  with  the  greatest  accumulation  of  carbonic  add 
in  the  blood-juices  and  tissues.  It  follows  that  the  nutritive 
processes  are  similar  in  both,  and  that  carbonic  acid  is  the 
principal  agent  for  producing  nutrition  in  animals.  Fur- 
thermore, it  is  the  most  soluble  of  all  the  organic  compounds 
of  carbon,  therefore  more  readily  diffused  through  the  tissues, 
while  little  force  is  needed  for  decomposing  it  in  the  metabolic 
processes  concerned  in  nutrition,  so  that  no  reason  presents 
why  this  so-called  waste  product  should  not  perform  an 
enormous  role  in  the  nutritive  processes,  or  the  same  as  in  the 
plant,  the  excess  passing  out  through  the  lungs  and  secretory 
functions  for  maintaining  a  balance  simply  ;  since  the  evolu- 
tion of  force  in  the  organism  by  producing  carbonic  acid 
would  naturally  create  an  excess  which  must  be  disposed  of. 
In  this  manner  nature  works  up  old,  effete  structures  in 
elaborating  new  tissues,  as  we  have  seen  her  do  in  the  case  of 
the  lymph  in  the  lymph-glands.  Why  not?  No  reason  on 
earth,  that  1  am  aware  of.  And,  being  utilitarian  to  the  last 
degree,  we  must  conclude  she  works  it  in  this  manner  with  car- 
bonic acid.  Last,  but  not  least,  the  deeply  suggestive  fact 
that  all  plant  and  animal  tissues  are  compounds  of  carbon, 
and  that  the  only  soluble  condition  of  carbon  fit  for  as- 
similation and  in  universal  distribution  over  the  earth  and 
in  living  organisms  is  carbonic  acid  gas.  And  why  interdict  in 
the  formative  processes  in  animals  ?  Do  not  seek  to  rob 
nature  of  her  most  available  means,  nor  place  too  wide  an 
interval  between  kindred  processes  in  plants  and  animals, 
seeing  that  they  are  only  grades  in  development  and  forms  of 
the  same  thing. 


CARBONIC   ACID   A   FOOD    FOR  THE   TISSUES.  367 

And  bear  in  mind,  also,  the  fact,  which  is  undeniable,  that 
increase  in  nutrition  corresponds  with  a  slow  circulation  and 
increase  of  carbonic  acid  in  the  blood  and  tissues.  This  cir- 
cumstance has  forcible  illustration  in  mania  apotu,  in  which 
sleep  has  at  last  been  induced  by  full  doses  of  morphia  or 
hydrate  of  chloral,  and  though  the  sleepless  and  terrified 
patient  is  thus  crushed,  so  to  speak,  into  sleep  at  the 
peril  of  his  life,  with  respiration  as  low  as  seven  and  eight 
per  minute,  breathing  stertorous,  the  skin  livid  from  ve- 
nous stasis  in  the  systemic  capillaries  (as  I  have  seen  in  a 
number  of  cases),  death  seeming  to  be  impending ;  yet  he  comes 
out  of  this  condition,  after  from  twelve  to  eighteen  hours'  sleep, 
actually  convalescent,  the  mind  clear  and  appetite  voracious. 
And  I  cannot  but  think  that  the  slow  respiration  and  circu- 
lation, together  with  the  great  accumulation  of  carbonic  acid 
in  the  system  which  must  inevitably  result,  are  the  chief 
elements  in  the  rapid  reparative  processes  in  these  cases.  The 
slowing  of  the  circulation  favoring  the  crystallizations  in  the 
nutritive  processes,  which  the  carbonic  acid  tends  to  increase ; 
at  the  same  time,  force  is  economized,  since  carbonic  acid  is 
necessarily  formed  in  maintaining  temperature  and  producing 
the  various  movements,  requiring  only  that  special  additions 
should  be  made  to  it — nitrogen,  hydrogen,  etc.  To  me  it  seems 
most  reasonable.  ' '  Eat  and  sleep' '  is  the  advice  in  the  nursery, 
and  the  child  most  gifted  in  this  respect  is  decidedly  the  finest. 
iStock  fatteners  limit  exercise  as  much  as  possible,  restricting 
the  animals  to  the  smallest  space,  with  the  object  of  making 
them  eat  and  sleep.  The  one  reduces  respiration  and  circu- 
lation ;  the  other  increases  the  nutritive  processes. 

And  as  age  creeps  on,  and  the  habits  become  more  and  more 
sedentary,  diminishing  respiration  and  circulation  in  propor- 
tion, for  the  same  reason  it  tends  to  the  accumulation  of  fat. 

Concerning  Carbonic  Acid  as  a  Stimulus. — The  burning 
sensation  in  the  throat  and  the  feeling  of  warmth  it  produces 
in  the  stomach  is  a  proof  that  carbonic  acid  acts  as  a  stimulus 
to  the  digestive  functions,  and  being  exceedingly  palatable,  is 
much  sought  after.  And  since  there  would  seem  to  be  enough 
and  to  spare  generated  in  the  system,  the  appetite  for  it  is 
very  probably  due  to  its  action  in  this  way.     In  impaired  di- 


308  CARBONIC  ACID  A   STIMULUS   TO   DIGESTION. 

gestion  it  is  highly  beneficial ;  and  irritable  stomachs  respond 
more  quickly  to  its  action  than  to  any  other  remedy,  espe- 
cially when  given  in  the  form  of  "mineral  water"  and  cham- 
pagne. 

But  perhaps  the  strongest  evidence  of  its  stimulating 
properties  is  furnished  by  its  action  upon  respiration,  being 
the  opposite  of  that  produced  by  oxygen,  which  acts  as  a 
sedative  to  the  respiratory  centre.  For  example,  if  pure  oxy- 
gen be  respired,  it  gradually  slows  respiration,  till  finally 
apncea  is  induced.  But  when  the  oxygen  is  discontinued  and 
carbonic  acid  substituted,  respiration  sets  in  again  at  once, 
growing  more  and  more  frequent ;  respiration  jumps  up  in 
leaps,  becoming  very  rapid  until  the  very  body  palpitates,  so 
there  can  be  no  doubt  that  it  is  a  respiratory  stimulant,  sub- 
serving useful  purpose  in  this  respect  for  maintaing  a  balance 
in  respiration.  And  oxygen  being  a  respiratory  sedative, 
the  necessity  for  a  special  stimulus  to  action  would  at  once 
appear  obvious,  while  this  increase  of  the  respiratory  rhythms, 
which  it  produces  when  in  excess,  soon  restores  the  dis- 
turbed equilibrium,  and  all  runs  on  as  before. 

In  similar  manner  it  also  affects  the  heart,  which  beats 
tumultuously  in  impending  asphyxia,  expanding  to  its  utmost 
limits  before  the  fatal  issue,  from  the  action  of  this  stimulus 
upon  the  nervous  centres  of  the  heart  intrinsic,  as  well  as 
extrinsic,  affecting  all  of  them,  and  the  dilator  equally  with  the 
contractor  nerves.  Indeed,  the  vaso-motor  centre  itself  is  in- 
cluded, as  is  fully  evidenced  in  the  wide  arc  of  movement 
described  by  the  so-called  Traube's  Curves  (Fig.  41),  which  is 
undoubtedly  produced  by  the  increasing  venosus  of  the  blood. 
There  can  be  no  doubt,  then,  that  carbonic  acid  is  a  stimulus 
to  the  respiratory,  circulatory  and  digestive  functions,  inclu- 
sive of  all  the  secretory  processes. 

One  of  the  most  difficult  circumstances,  however,  to  account 
for  occurs  in  connection  with  sleep,  when  respiration  and  circu- 
lation are  slower  than  at  any  other  time — this  notwithstanding 
the  fact  of  an  increase  of  carbonic  acid  in  the  blood.  This, 
however,  could  be  accounted  for  by  the  fact  that  there  is 
similar  obtundity  in  all  of  the  nervous  centres  ;  sight,  hear- 
ing, smelling,  tasting,  feeling — all  are  obtunded ;    so  that 


PHYSIOLOGY    OF   SLEEP.  369 

unusual  force  must  be  applied  to  either  one  of  them  in  order 
to  increase  the  action  :  not  produced  by  brain  anaemia,  either, 
only,  in  one  sense,  that  there  is  diminished  flow  of  arterial 
blood  in  the  brain,  since  the  veins  and  capillaries  are  liter- 
ally distended  with  venous  blood,  and  the  brain  larger  in 
consequence,  occupying  more  room  than  in  the  waking  con- 
dition ;  which  undoubtedly  favors  the  nutritive  processes, 
having  this  as  the  end.  But,  then,  what  induces  it?  The 
necessity  for  repair  !  The  arteries  contract  and  diminish  the 
lumen  in  the  vessels,  while  the  slowing  of  respiration  causes 
the  venous  blood  to  dam  back  in  the  sinuses  and  cerebral 
veins,  and  so  producing  it,  that  the  nutritive  processes  may 
restore  what  is  lost  by  attrition  in  evolving  force  or  for  pro- 
ducing the  movements  in  animal  life.  Very  well ;  we  must 
accept  that !  Cause  of  causes  ! — show  us  the  rest  of  it !  Fall- 
ing back  upon  the  inevitable  metamorphosis  of  force,  we  find 
that  the  arterial  blood  ebbs  and  flows  in  the  brain  according  to 
whether  sleep  or  animation  is  desirable,  nutrition  or  force 
most  needed  in  the  organ  and  organism  ;  but  there  we  stop  at 
the  brink  of  the  abyss  in  Force  itself,  involving  the  universe 
of  matter,  for  matter  and  force  are  forms  of  the  same  thing, 
undoubtedly. 

Concerning  the  Functions  of  Nitrogen  Gas  in  the  Animal 
Organism. — Briefly,  an  amount  of  nitrogen  disappears  in  res- 
piration to  reappear  in  arterial  blood,  but  again  to  disappear  in 
the  tissues,  since  the  venous  contains  less  than  arterial  blood, 
the  relative  proportion  being  1.30  to  1.60  in  the  100.  But 
when  we  come  to  the  gases  contained  in  its  several  portions  of 
the  intestinal  canal,  this  phenomenon  has  ready  explanation, 
constituting  a  necessary  part  of  the  mechanics  in  the  diges- 
tive and  absorptive  processes  in  the  organs,  or  means  to  ends, 
the  secretion  of  the  gas  being  essential  to  the  maintenance  of 
a  balance  in  pressure  within  the  canal,  which,  of  course,  must 
have  adjustment  with  the  exigencies  in  the  functions,  and  as 
a  consequence  must  vary  in  the  several  portions.  Thus,  in  a 
series  of  experiments  instituted  upon  executed  criminals,  by 
Majendie  and  Chevreul,  the  following  gases  were  found  to  be 
present  in  the  stomach  and  intestines  : 


370 


FUNCTIONS    OF    NITKOGFJ5T   GAS. 


Gases  contained  in  the  Stomach. 

Oxygen 11.00 

Carbonic  acid 14.00 

Pure  hydrogen 3.55 

Nitrogen 71.45 

100.00 

Thus,  casting  our  eye  over  the  chemical  analyses,  we  find 
that  oxygen  in  the  proportion  of  11  parts  in  the  100  presents  for 
the  first  and  last  time  in  the  stomach  ;  hence,  is  very  probably 
carried  in  with  the  boluses  and  liquids,  whence  it  is  absorbed 
and  carried  to  the  liver,  to  be  consumed  in  the  metabolic  pro- 
cesses ;  a  remnant,  as  it  were,  of  the  primitive  mode  of  respi- 
ration. It  would  also  account  for  a  portion  of  the  nitrogen 
and  carbonic  acid,  but  not  all  of  them,  leaving  a  large  residuum 
still  unaccounted  for,  save  by  the  secretory  function  in  the 
gastric  capillaries.  This  circumstance  is  fully  proven  in  the 
cases  of  the  small  and  large  intestines,  notably : 


Oases  contained  in  the  Small  Intestine. 

First  Second 

criminal.  criminal. 

Carbonicacid 24.39  40.00 

Pure  hydrogen 55.53  51.15 

Nitrogen 20.08  8.85 

100.00 


Third 

criminal. 

25.00 

8.40 

66.60 


100.00 


100.00 


Gases  contained  in  the  Large  Intestine. 


First  Second 

criminal.  criminal. 

Carbonic  acid 43.50  70.00 

Carbureted  hydrogen  and  traces 

of  sulphureted  hydrogen 5.47  

Pure    hydrogen   and  carbureted 

hydrogen 11.60 

Pure  hydrogen 

Carbureted  hydrogen .... 

Nitrogen : 51.03  18.40 


Third 
criminal. 
Caecum. 
12.50 


Third 
criminal. 
Rectum. 
42.86 


7.50 
12.50 
67.50 


11.18 


45.96 


100.00  100.00  100.00  100.00 

It  will  be  seen  from  the  above  exhibit  that  carbonic  acid 
and  nitrogen  are  the  principal  gases  in  the  small  and  large  in- 
testines, as  well  as  in  the  stomach;  in  the  third  criminal  the  total 
amount  aggregating  as  much  as  91  and  88.  ^2  respectively  for 
the  small  and  large  intestines,  the  remainder  being  made  up  of 
the  hydrogen  gases,  principally  pure  hydrogen. 

The  large  amount  of  this  gas  in  the  small  intestines  of  the 


FUNCTIONS    OF    NITROGEN   GiS.  371 

first  and  second  criminals  is  difficult  of  explanation ;  but  as 
hydrogen  is  evolved  in  the  nutritive  processes  in  the  plant,  we 
may  infer  that  the  epithelial  cells  are  the  principal  source.  But 
there  can  be  no  doubt  respecting  the  source  of  the  remarkable 
quantity  of  carbonic  acid  and  nitrogen  in  the  large  intestine 
of  all  the  criminals,  being  respectively  94.53.  88.40,  80  and 
88.82  in  the  rectum  of  the  third  criminal ;  the  quantities  of  the 
two  gases  varying. 

All  we  seek  to  show  by  this  circumstance  is  the  power  on  the 
part  of  the  blood-capillaries  to  secrete  these  gases  out  of  the 
blood  in  the  measure  of  the  physiological  requirements  in  the 
organs  in  the  exigencies  in  the  functions  ;  and  since  the  whole 
matter  relates  to  pressure,  the  relative  amount,  of  course, 
would  have  to  be  determined  by  the  quantities  of  the  other 
gases  already  in  the  intestines  or  evolved  by  the  chemical  and 
secretory  processes  in  the  organs  ;  and  as  this  must  necessarily 
vary  from  time  to  time,  it  would  imply  a  corresponding  power 
of  absorbing  them  when  in  excess  of  the  demand  ;  hence  the 
circumstance  of  the  excess  in  expiration  daring  digestion,  as 
has  already  been  mentioned. 

Finally,  the  power  on  the  part  ol  the  intestines  to  secrete 
the  gases  is  proven  to  demonstration  by  physiological  experi- 
ment upon  the  gut  (p.  203),  while  respiration  itself  is  based 
upon  this  power  in  the  blood-vessels  to  secrete  and  absorb  the 
gases  ;  otherwise  carbonic  acid  could  not  be  excreted  nor 
oxygen  and  nitrogen  be  absorbed. 

In  Reference  to  the  Odor  of  F&cal  Matter. — In  respect 
to  the  peculiar  odor  of  faecal  matter,  the  chemical  experi- 
ments of  Professor  Liebig  afford  the  clue.  For  example,  he 
ascertained  that  if  albuminous  compounds  are  subjected  to 
heat  with  solid  hydrate  of  potash,  and  the  heat  be  continued 
until  the  greater  portion  or  the  whole  of  the  nitrogen  is  dis- 
sipated as  ammonia,  and  free  hydrogen  begins  to  escape,  the 
residue,  when  supersaturated  with  dilute  sulphuric  acid,  and 
distilled,  yields  a  liquid  containing  acetic  and  butyric  acid,  and 
possessing  in  a  very  intense  degree  the  peculiar  and  char- 
acteristic odor  of  human  fasces;  the  odor  varying  accord- 
ing to  the  substance  used,  in  this  way  accounting  for  all 
varieties  of  faecal  smell.     Now,  then,  in  view  of  the  fact  that 


372  PHYSIOLOGY   OF   FJEOAL    ODOR. 

the  chemical  reactions  in  living  organisms  without  the  agency 
of  heat  per  se,  but  by  the  action  of  electrical  force,  aided  by 
the  mutual  affinities  in  the  molecules,  and  that  oxygen  and 
lrydrogen  in  the  form  of  water  have  their  bond  of  union 
Droken  in  this  manner,  notwithstanding  the  prodigious  force 
it  involves,  the  explanation  of  the  chemical  reactions  detailed 
by  Professor  Liebigfor  evolving  fa9cal  odor  without  involving 
any  harm  to  the  tissues,  would  appear  natural  enough,  and  not 
at  all  extraordinary. 

Concerning  the  Genesis  and  Functions  of  the  Gases  in 
the  Air -Bladders  of  the  Fishes. — This  function  in  the  blood- 
vessels for  carrying  the  gases  to  and  from  the  internal  parts 
for  subserving  important  mechanical  uses  in  connection  with 
the  special  functions  in  the  organs  has  forcible  illustration  in 
the  air-bladders  of  the  fishes,  in  which  it  subserves  a  dual 
function,  notably : 

1.  For  buoying  them  in  the  media,  thereby  powerfully 
assisting  the  action  of  the  fins,  enabling  the  animal  to  ascend 
or  descend  with  the  utmost  ease  and  celerity  by  simply 
expanding  and  contracting  the  air-bladders  and  body-walls, 
using  the  fins  and  tail,  of  course,  for  assisting  the  action 

2.  For  transmitting  sonorous  vibrations  through  the  body- 
walls  upon  the  ossicles  and  otolites  of  the  auditory  apparatus 
(Fig.  147,  o,  m,  I,  d).  As  will  be  seen,  the  anterior  portions  of 
the  bladder  (p)  fit  accurately  against  the  expanded  base  of  the 
large  ossicle  (o),  this  against  the  ossicles  m  and  I,  whence  the 
force  is  transmitted  upon  the  delicate  extremities  of  the  acoustic 
nerve,  which  are  expanded  upon  the  chamber  of  the  vestibule 
Id j  means  of  the  two  subspherical  "atria"  on  the  body  of 
the  atlas,  close  to  the  foramen  magnum,  and  the  endolymph 
which  fills  both  atria  and  the  common  sinus.  The  sonorous 
vibrations  thus  communicated  to  the  suspended  otolites  (d)  are 
made  to  beat  upon  the  nervous  filaments  of  the  auditory  nerve 
in  the  chamber  of  the  vestibule  containing  the  otolites  upon 
the  lining  membrane  of  which  the  nerves  are  expanded. 

In  the  herring,  the  tubular  prolongation  of  the  fore  part  of 
the  bladder  (Fig.  148,  Jc)  advances  to  the  basi-occipital  and 
bifurcates  ;  each  branch  penetrates  the  side  of  the  base  of  the 
skull,  again  bifurcates,  and  terminates  in  two  blind  sacs,  which 


FUNCTIONS  OF  THE  GASES  IN  FISH-BLADDEKS. 


373 


374  FUNCTIONS  OF  THE  GASES  IN  FISH-BLADDERS. 

are  in  contact  with  similar  csecal  processes  of  the  labyrinth 
(Owen). 

In  this  manner,  then,  the  gases  contained  in  the  air-bladders 
of  the  fishes  subserve  two  important  functions  :  one  for  over- 
coming body-inertia,  the  other  for  transmitting  sonorous  undu- 
lations upon  the  auditory  nerves  ;  while  in  the  mammalia,  we 
have  seen  that  air  is  utilized  in  the  intestines  for  increasing  the 
digestive  and  absorptive  processes  by  means  of  the  force  trans- 
mitted from  the  walls  of  the  gut  and  the  muscular  envelope  of 
the  body  upon  the  aliment ;  or,  -in  other  words,  for  overcoming 
inertia  in  the  nutritive  and  force-producing  elements  and  speed- 
ing them  into  the  vascular  channels,  and,  in  very  truth,  is  the 
stepping-stone  in  higher  development  or  mammalian  evolution. 
And  that  the  blood  is  the  source  of  the  gases  in  the  air- 
bladders  of  the  fishes  is  manifest,  from  the  following  facts, 
notably :  1.  The  air-bladder  is  the  nascent  condition  of  the 
lungs,  into  which  it  is  subsequently  developed ;  hence  the 
power  to  excrete  and  absorb  the  gases  inheres  in  the  organ. 

2.  It  would  account  for  the  absence  of  the  air-duct  in  vast 
numbers  of  fishes,  notably,  Acanthopteri,  e.  g.,  perch,  mullet, 
mackerel,  angler,  etc.;  Anacanthini,  e.  g.,  cod!  plaice,  etc.  ; 
Plectognathi,  Lophobranchi.  And  there  being  no  air-duct  in 
these  fishes,  it  follows  that  the  blood  must  be  the  source  of  the 
gases  contained  in  the  air-bladder,  and  not  the  intestinal  canal, 
which  serves  as  an  outlet  in  the  other  fishes  for  rapid  discharge 
of  the  gases,  in  order  to  obviate  strain  when  moving  from  the 
deep  to  the  surface  waters,  where  sudden  expansion  is  inevita- 
ble. 

3.  It  would  account  for  the  special  gases  in  the  air-bladders 
of  the  fishes,  which  consist,  in  most  fresh-water  fishes,  of  nitro- 
gen and  a  very  small  quantity  of  oxygen,  with  a  trace  of  car- 
bonic acid  gas ;  while  in  the  air-bladder  of  sea-fishes,  espe- 
cially those  which  frequent  great  depths,  oxygen  predominates, 
as  much  as  1-7  in  the  100  in  volume  being  reported  by  Biot  in 
some  of  the  deep-sea  Mediterranean  fishes ;  the  rest  nitrogen, 
with  a  trace  of  carbonic  acid.  In  these  cases,  the  air-bladders 
function  as  special  reservoirs  for  oxygen  to  subserve  respira- 
tory purposes  in  the  depths  where  the  gas  does  not  circulate 
so  freely  ;  finally,  no  hydrogen  has  ever  been  detected  in  the 


GENESIS    OF   THE    GASES    IN   FISH-BLADDEKS.  375 

air-bladders  of  fishes,  which  also  shows  the  gases  are  not  de- 
rived from  the  intestinal  canal. 

4  Lastly,  we  have  to  mention  the  peculiarities  that  obtain 
in  the  capillary  network  of  the  air-bladders  as  the  special  pro- 
vision for  excreting  and  absorbing  the  gases,  while  the  branchiae 
form  an  open  portal  for  their  ingress  and  egress  to  the  blood 
vessels.  The  principal  seat  of  the  vascular  ramifications  in  the 
air-bladder,  like  that  in  a  true  lung,  is  the  mucous  lining  mem- 
brane, but  there  is  variety  in  the  terminal  divisions  of  the 
arteries.  In  the  carp,  for  example,  they  terminate  in  fan-like 
tufts  over  almost  every  part  of  the  inner  surface.  In  the  pike 
they  are  larger  and  more  localized,  but  without  any  special 
aggregation  of  the  capillaries  to  form  a  "  vaso-ganglion" ;  but  in 
the  perch  and  cod  the  capillaries  are  aggregated  so  as  to  form 
red,  gland-like  bodies  (Fig.  149) ;  the  capillaries  reuniting 
into  larger  vessels,  which  again  ramify  around  the  gland-like 
body  ;  the  rest  of  the  inner  surface  of  the  air-bladder  retains 
the  ordinary  simple  capillary  system. 

It  will  be  seen  that  the  afferent  and  efferent  vessels  to  these 
bodies  form  vascular  loops,  which  are  covered  by  a  layer  of 
vessels  and  epithelium  (a,  a).  In  addition  to  this,  however, 
are  a  number  of  peculiarly  arranged,  elongated  corpuscles, 
which  depend  in  two  rows  from  each  vascular  branch,  and  are 
bound  together  by  a  loose  cellular  tissue  ;  the  corpuscles  are 
beset  with  fine  villiform  processes.  The  blood  returns  from 
the  vaso-ganglions  by  small  veins,  which  rarely  accompany — 
more  commonly  cross — the  arteries  (Owen),  and  is  certainly 
strong  corroborative  proof  of  this  function  in  the  arterial 
capillaries  for  excreting  the  gases.  In  the  eel  and  conger,  the 
two  chief  ganglions,  which  are  situated  at  the  sides  of  the 
opening  of  the  air-duct,  consist  of  both  arterioles  and  venules  ; 
they  consist  of  straight  parallel  capillaries  (Fig.  150) ;  their 
efferent  trunks  do  not  ramify  in  the  immediate  margin  of  the 
vaso-ganglion  from  which  they  issue,  as  in  the  vaso-ganglions 
of  the  cod,  burbot,  acerine  and  perch,  but  run  for  some  distance 
before  they  again  branch  to  form  the  common  capillary  system 
of  the  lining  membrane  of  the  air-bladder  (Owen). 

This  aggregation  of  capillaries  in  arterial  and  venous  retia  is 
undoubtedly  favorable  for  rapid  secretion  and  absorption  of 


376 


GKNESIS    OF   THE   GAS'S    IN    FISH-BLADDEKS. 


the  gases ;  indeed,  not  gases  only,  but  liquids  as  well,  since 
the  Malpighian  glomeruli  are  homologous,  formed  by  the 
terminal  branches  of  the  renal  artery,  and  from  which  an 
afferent  vessel  is  given  off  to  be  distributed  to  the  urinary 
tubules  (Fig.  117,  va,  gl,  ve).     And  they  also  occur  in  other 


Fig.  148. — Abdominal  Viscera,  with  Air-Bladder  (k)  in  situ,  Herring  (reduced).— Brandt 
and  Ratzeburg.  .  Medizinische  Zoologie.  4to.  1833.  a,  CEsophagus  ;  b,  stomach  ; 
c,  pylorus  ;  d,  appendices  pyloricae  ;  e,  e,  intestine  ;  /,  anus  ;  7i,  h,  testes  ;  i,  genital 
ducts  ;  k,  air-bladder  ;  I,  pneumatic  duct. 


Fig.   149. Superficial  and  Looped  Vessels  of  the  Vaso-Ganglion  of  the  Air-Bladder, 

Cod.— Dr.  Wdliams. 


Fig.  150.— Parallel  Vessels  of  the  Vaso-Ganglion  of  the  Air-Bladder,  Eel.— Dr.  Williams. 

localities — notably,  in  the  arteries  of  the  mesentery  contiguous 
to  the  intestines — and  are  common  in  the  limbs  of  the  sloth, 
the  axillary  and  iliac  arteries  in  these  slow-moving  animals, 
just  before  entering  the  limbs,  suddenly  dividing  into  numer- 
ous small  channels,  which  again  unite  into  one  trunk  before 
the  members  are  given  off ;  in  these  cases  serving  as  reser- 
voirs for  storing  arterial  blood  for  evolving  the  local  actions 


OFFICE  OF  RhTE  MIR  A  BILE.  377 

concerned  in  digestion,  and  for  producing  the  muscular  force 
for  sustaining  the  weight  of  the  animal  suspended  by  its  limbs. 
But  this  is  seen  upon  a  prodigious  scale  in  the  enormous  arte- 
rial plexuses  in  Cetacea,  wherein  a  vast  quantity  of  arterial 
blood  may  be  accumulated  for  supplying  the  wants  of  the 
animal  during  prolonged  periods  of  submersion.  For  exam- 
ple, the  intercostal  arteries  divide  into  a  vast  number  of 
branches,  which  run  in  a  serpentine  course  between  the  pleura 
and  the  ribs,  and  penetrate  the  intercostal  muscles,  every- 
where lining  the  walls  of  the  thorax ;  moreover,  they  pass 
in  between  the  ribs  near  their  articulation,  and  anastomose 
extensively  with  each  other.  And  in  like  manner  the  central 
nervous  system  is  embossed  by  dense  arterial  plexuses  ;  more 
especially  the  medulla  oblongata,  where  a  thick  substance  is 
formed  by  their  ramiii cations  and  convolutions — aretemira- 
bile  on  a  prodigious  scale  for  sustaining  the  functions  in  the 
great  nervous  centre  for  the  organism  during  prolonged  periods 
of  submergence,  Nature  in  this  manner  storing  oxygen  for  the 
purpose.     Mirabile ! 


CHAPTER  XVII. 

FACTS  IN  DEVELOPMENT,  SHOWING  THE  RELATIONS  WniCH 
THE  HE^RT  SUSTAINS  TO  THE  MECHANICS  OF  CIRCULA- 
TION. 

Principle  in  Cardiac  Evolution— Respiration  and  Circulation  in  "Worms — The  First  Indi- 
cations of  a  Heart  Found  in  Connection  with  the  Localization  of  the  Respiratory 
Apparatus,  notably  Terebella — Facts  Revealed  in  Decapods — Ditto  Fishes — Ditto 
Reptiles — Differentiation  of  the  Left  Cardiac  Chambers  with  Air-Breathers,  First 
the  Auricle,  then  Ventricle — Enormous  Development  of  the  Muscles  in  the  Heart 
in  Tortoise — Reasons  Therefor — Perfection  of  the  Interventricular  Septum  in 
Crocodilia,  but  Leaving  a  Passage  between  the  Left  and  Right  Sides  of  the  Heart,  so 
as  to  Allow  Reflux  during  Submergence — Reasons  Therefor — The  Changes  with 
Progress  in  Development  till  the  Birds  are  Reached,  in  which  Cardiac  Develop- 
ment is  Complete — The  Heart  always  in  Intimate  Connection  with  the  Oxygenating 
Apparatus,  since  it  Relates  to  the  Evolution  of  Force  in  the  Organism. 

In  tracing  cardiac  development,  we  must  begin,  of  course, 
with  the  principle  upon  which  development,  itself,  is  based, 
when  all  the  phenomena  appertaining  to  it  have  ready  explana- 
tion ;  otherwise  are  inexplicable.  Since  all  the  activities,  in- 
clusive of  the  movements  in  the  viscera,  blood-vessels,  and 
heart — indeed,  every  variety  of  motion  in  the  organism — 
is  evolved  from  force  which  is  generated  by  the  combinations 
effected  with  oxygen,  it  follows  that  the  importation  and 
circulation  of  oxygen  is  fundamental  in  the  organism ;  and 
the  blood  being  the  medium  for  receiving  and  transmitting  it 
to  the  tissues,  the  comprehensive  arrangements  that  obtain  for 
pumping  it  through  the  lungs  and  tissues  in  the  measure  of 
the  physiological  requirements  will  be  readily  apprehended  ; 
also,  why  the  respiratory  and  circulatory  apparatus  should  be 
a  connected  movement,  otherwise  the  scheme  would  fail  The 
intimate  relations  which  the  heart  sustains  to  the  respiratory 
apparatus,  then,  are  to  be  explained  by  this  principle  in  the 
mechanics ;  while  the  changes  of  form  which  it  undergoes 
with  progress  in  development,  are  due   to  the  peculiarities 


PULSATION  FIEST  ITST   THE  VESSELS.  379 

which  obtain  in  the  respiratory  apparatus,  which  impress 
themselves  upon  the  heart  as  the  local  reservoir  and  force- 
pump,  for  receiving  and  transmitting  the  blood  to  and  from 
these  organs,  together  with  the  necessity  for  a  gradual  and 
more  and  more  complete  separation  of  the  arterialrzed  blood, 
upon  which  progress  in  development  depends,  since  this  is 
essential  for  the  production  of  the  relative  phenomena. 

In  fine,  the  vascular  system  is  molded,  so  to  speak,  to  the 
respiratory  apparatus,  undergoing  special  modifications  in  cor- 
respondence with  the  changes  which  have  taken  place  in 
the  stage  in  development,  while  the  differentiation  of  a  heart 
or  central  force-pump  in  connection  with  the  oxygenating 
apparatus  is  for  the  purpose  of  increasing  circulation  in  it  in 
correspondence  with  the  amount  of  force  which  is  expended  in 
the  organism,  the  one  involving  the  other.  And  viewed  from 
this  stand-point  and  the  law  for  increasing  circulation — namely, 
by  rhythmical  changes  in  pressure — the  special  role  in  the  heart 
and  the  mechanical  principle  it  involves,  inclusive  of  the  nerves 
for  coordinating  it  with  respiration,  is  at  once  made  intelligible, 
together  with  all  the  relative  phenomena  appertaining  to  it ; 
otherwise  inexplicable.  Thanks  to  numerous  workers  in  the 
field  and  the  opulence  of  material,  the  labor  involved  in  this 
inquiry  is  enormously  diminished,  requiring  only  to  be  care- 
fully collated  and  systematically  arranged.  For  this  purpose, 
we  begin  with  the  earliest  stages  in  cardiac  development ; 
notably,  worms,  in  which  a  highly  complex  circulation  is  car- 
ried on  in  the  absence  of  a  heart  for  producing  it,  following 
thence  through  the  successive  stages  in  cardiac  evolution  to 
the  warm-blooded  animals,  in  which  cardiac  development  is 
complete.  And  we  begin  with  the  worms,  for  the  reason 
that  pulsation  is  first  visible  in  the  vessels ;  hence,  is  not 
necessarily  dependent  upon  the  heart  or  synonymous  with  its 
action,  since  it  exists  long  before  a  heart  comes  into  the  scheme  ; 
moreover,  is  not  limited  to  the  blood- vascular  system,  for  it  is 
met  with  in  the  veins  and  lymphatics  of  amphibia  and  occurs  in 
the  lacteals  of  warm-blooded  animals.  At  the  same  time,  how- 
ever, in  the  worms  local  dilatations  occur  in  the  vessels  corre- 
sponding with  the  branchial  and  air-vesicles,  thus  early  indi- 
cating the  principle  in  the  blood- vascular  system  for  increasing 


380 


GROUND    PLAN    OF   THE   CIRCULATION. 


circulation  in  the  lungs  by  the  differentiation  of  a  heart  for 
assisting  the  action. 

Briefly  summarized,  the  ground  plan  of  the  animal  circu- 
lation commences  by  the  formation  of  several  longitudinal 
vessels  (three  or  more),  extending  the  length  of  the  body  of 
the  worms,  first  visible  in  Nemertina  (Gegenbaur) ;  a  median 
dorsal,  which  lies  above  the  enteron  and  is  pulsatile,  and  two 


Fig.  151. — Ground  Plan  of  the  Vascular  System  in  Nemertina. — Gegenbaur.  d,  Dorsal 
longitudinal  trunk  ;  I,  I,  lateral  longitudinal  vessels.  The  arrows  indicate  the  direc- 
tion of  the  stream. 

lateral  vessels,  one  upon  either  side  (Fig.  151,  d,  I,  Z),  which  are 
veins  that  deliver  the  blood  into  the  median  dorsal  artery,  with 
which  they  connect  at  the  terminal  ends  and  by  transverse 
vessels  ;  whence  it  is  passed  forward  by  successive  expansions 
and  contractions  in  the  dorsal  vessel,  the  lateral  branches 
given  off  at  regular  intervals  by  this  great  trunk  conveying  it 


NO   HEART    IN"    THK   WORMS. 


3  SI 


to  the  muscular  parietes  for  generating  force,  and  to  the  cuta- 
neous capillaries  for  oxygenation,  to  be  re-collected  in  the  cor- 
responding branches  of  the  lateral  veins,  which  also  collect 
the  blood  in  the  viscera  ;  while  a  portion  continues  on  to  the 
cephalic  end  and  the  vascular  loop  or  loops,  as  the  case  may 
be,  which  encompass  the  oesophagus.  The  simplest  condition 
of  the  vascular  system  is  seen  in  those  which  have  no  per- 
ceptible respiratory  organs,  oxygenation  being  effected  through 
the  thin  integument  and  subjacent  capillary  network — 
e.  g.,  nais  and  planar  ia.  In  the  transparent  body  of  erpobdella 
vulgaris,  two  median  vessels,  a  dorsal  and  a  ventral  vessel,  are 
discernible,  which,  by  their  waves  of  pulsation,  convey  the 
blood  into  numerous  lateral  branches  (Fig.  152,  5),  the  capil- 

Fig.  153.  Fig.  153. 


Fig.  152.—  Diagrammatic  Representation  of  the  Vascular  System  in  Erpobdella  Vulgaris, 
showing  the  numerous  lateral  branches  given  off  by  the  median  ventral  vein  (b),  and 
which  anastomose  with  corresponding  branches  from  the  two  lateral  venous  trunks 
(c,  c),  and  dorsal  artery  (a). 

Fig.  153. — Vascular  System  of  Lumbricus  Terrestris  (anterior  portion),  a,  a,  Median 
ventral  vein  ;  6,  b,  great  dorsal  artery  ;  c,  c,  sacculated  arches.  The  arrows  show 
direction  of  the  stream. — Grant. 

laries  of  which  are  continuous  with  those  of  the  two  great 
returning  veins  (c,  c),  extending  backward  along  the  sides  of 
the  body  (Morren).  In  annelides,  it  is  pretty  much  the  same ; 
the  venous  blood  of  these  animals  is  commonly  returned  from 
the  system  to  the  posterior  extremity  of  the  dorsal  artery  by 
a  median  ventral  vein,  or  by  two  inferior  lateral  veins  ;  but 
considerable  modifications  are  induced  in  higher  genera  by 
development  of  respiratory  organs  in  the  form  of  external 
cephalic  or  dorsal  branchiae  or  internal  air- vesicles.  In  nereis 
cuprea,  the  long  dorsal  artery  appears  slightly  dilated  in  each 


382  NO   HEART  IN    THE   WORMS. 

segment  of  the  body,  and  receives  or  gives  off  the  branchial 
vessels  from  the  arterial  arches  which  encompass  the  oesopha- 
gus (Grant)  In  other  species,  the  branchial  vessels  are  given 
off  to  these  organs  from  each  side  of  the  dorsal  artery  in  its 
whole  course  forward  (Fig.  153,  d,  b,  b),  and  the  fifteen  pairs  of 
ramose  branchiae  presenting  in  arenicola  induce  as  many  corre- 
sponding modifications  in  the  vascular  system  of  these  ani- 
mals ;  while  small  pulsating  vesicles  are  generally  perceptible 
on  the  lateral  systemic  branches  of  the  dorsal  artery. 

In  the  earth-worm  {Jumbricus  terrestris).  the  direction  of  the 
internal  currents,  owing  to  the  greater  transparency  of  the 
body,  is  more  perceptible  than  in  the  more  opaque  body  of 
the  leech,  where  it  is  necessary  to  examine  this  part  in  very 
young  individuals,  and  where  the  currents  have  appeared 
often  to  change  their  direction  through  the  vascular  trunks. 
Successive  waves  of  contraction  are  distinctly  seen  in  the 
earth-worm,  extending  from  behind  forward  along  the  wide 
dorsal  vessel ;  and  by  removing  the  integuments  and  pressing 
this  artery  between  the  forceps,  it  becomes  empty  in  front  and 
turgid  behind.  It  appears  to  receive  the  arterialized  blood 
from  the  air-vesicles,  and  sends  off  numerous  lateral  branches 
in  its  course,  especially  to  the  alimentary  canal  and  the  geni- 
tal organs. 

The  venous  blood  is  collected  from  the  viscera  chiefly  by 
the  great  median  subgastric  or  epineural  vein,  extending 
backward  between  the  digestive  canal  and  the  nervous  columns; 
and  this  vessel  appears  to  send  off  branches  to  the  numerous 
minute  respiratory  vesicles.  A  small  inferior  median  vessel  or 
hyponeural  vein  is  also  perceived,  extending  along  the  under 
surface  of  the  nervous  chords,  and  an  accompanying  lateral 
branch  is  seen,  as  usual,  on  both  sides  of  the  same  columns. 

Anterior  to  the  commencement  of  the  stomach,  the  great 
dorsal  artery  (Fig.  153,  b,  b)  communicates  with  the  median 
subgastric  vein  (a,  a)  by  five  or  more  pairs  of  lateral,  wide,  sac- 
culated arches  (c,  d),  which  embrace  the  oesophagus,  as  the 
corresponding  vascular  arches  which  connect  these  t  wo  vessels 
in  other  annelides  and  in  the  entomoid  classes.  According 
to  Dr.  Williams,  the  following  is  the  plan  of  circulation  in 
earth-worms  (Fig.  154) :    The  circulation  is  very  complicate. 


NO   HEART    IN   THE   LEECH. 


383 


They  respire  through  the  cutaneous  surface,  which  is  occu- 
pied by  a  dense  capillary  plexus  ;  also  through  the  alimentary 
canal,  which  is  similarly  supplied,  and  which  undoubtedly 
functions  as  a  respiratory  as  well  as  digestive  organ.  It  is  also 
claimed  that  the  stratum  of  viscid  matter  in  which  they  are 
always  enveloped  is  remarkably  endowed  with  the  property  of 
absorbing  and  dissolving  atmospheric  air  (Jones). 


Eig.  154. — Diagram.  Illustrative  of  the  Circulatory  Apparatus  in  the  Leech  (Hirudo 
Medici >. alis). — After  Dr.  Williams,  a,  Great  dorsal  vessel  ;  c,  ventral  vessel ;  d,  d, 
intercommunicating  vessels  between  dorsal  and  ventral  trunks  ;  e,  e,  lateral  abdomi- 
nal trunks  ;  g,  vessels  distributed  over  the  cascal  appendages  to  the  stomach. 

From  the  equal  development  of  the  great  vascular  trunks 
and  the  numerous  transverse  anastomoses,  it  is  easy  to  per- 
ceive how  the  circulation  in  these  animals  can,  by  the  closing 
of  the  divided  ends  of  the  vessels,  become  accommodated  to 
extensive  mutilations,  and  proceed  without  interruption  in  a 
few  segments  detached  from  the  trunk.  Some  of  the  simpler 
forms  of  annelides,  e.  g.,  styluria,  are  thus  enabled  to  ex- 


384  NO   HEART  IN-  THE   LEECH. 

tend  their  means  of  propagation  by  the  spontaneous  transverse 
fission  of  the  body. 

But  the  leech,  perhaps,  presents  the  most  complicate  vascular 
arrangements  of  any  of  the  annelides,  though  the  ground  plan 
of  the  circulation  is  fundamentally  the  same  as  in  all  the  other 
worms  and  entomoid  classes,  consisting  of  a  number  of  longi- 
tudinal trunks  with  the  median  dorsal  and  ventral  occupying 
the  old  positions,  the  blood  passing  to  the  caudal  end  through 
the  one  and  the  cephalic  in  the  other,  while  the  lateral  branches 
pass  into  the  segments  and  effect  circulation  transversely. 
The  dorsal  is  recipient  of  arterialized  blood,  which  it  dis- 
tributes to  the  organs  (but  venous  blood  also  flowing  into  it 
by  the  anastomosing  branches,  especially  with  the  median 
ventral),  while  the  ventral  functions  as  the  systemic  venous 
trunk,  from  which  the  blood  is  sent  to  the  respiratory  vesicles. 
In  addition  to  the  dorsal  and  ventral  trunks,  however,  are  two 
large  lateral  trunks,  one  on  each  side  (Fig.  155,  e,  e),  which, 
according  to  M.  Duges,  *  are  appropriated  to  the  respiratory 
system  of  lateral  sacculi,  of  which  there  are  seventeen  pairs 
symmetrically  disposed  along  the  sides  of  the  body,  with  thin 
spiracles  opening  along  the  ventral  surface. 

That  the  movement  of  the  blood  in  the  lateral  or  respi- 
ratory system  of  vessels  is  quite  distinct  from  that  which  is 
accomplished  in  the  dorsal  and  ventral  or  systemic  trunks ; 
sometimes  it  passes  down  one  of  these  vessels  from  the  head 
toward  the  tail,  and  in  an  opposite  direction  on  the  other  side 
of  the  body  ;  but  in  a  short  time  the  movement  of  the  cur- 
rents will  be  seen  to  become  completely  reversed,  so  that  an 
undulatory  motion,  rather  than  a  complete  circulation,  is  kept 
up.  By  this  action  of  the  lateral  canals  the  blood  is  made 
perpetually  to  pass  and  repass  the  respiratory  sacculi  ;  and, 
opposite  to  each  of  these,  branches  are  given  off  which  form 
so  many  independent  vascular  circles,  representing  very  closely 
the  minor  or  pulmonary  circulation  of  higher  animals. 

Finally,  the  rich  supply  of  arterialized  blood  sent  to  the 
lateral  caeca  of  the  stomachal  cavity  for  promoting  the  secre- 
tory processes  in  these  organs  in  connection  with  digestion,  has 

*  Ann.  des  Sci.  Nat.,  vol.  xv. 


NO   HEAET   I  INT   THE   LEECH. 


385 


forcible  illustration  in  the  beautiful  arborescent  arrangement 
of  the  vessels  in  the  walls,  with  their  footstalks  connecting 
immediately  with  the  dorsal  artery  (Fig.  155,  g,  a).  The  fol- 
lowing illustration  (Pig.  156)  will  give  some  idea  of  the  rela- 
tions the  csecal  pouches  ( 7c)  sustain  to  the  stomachal  cavity 
and  respiratory  vesicles  ;  the  stomach  itself  occupying  about 
two-thirds  of  the  visceral  cavity,  is  divided  by  septa  or  dia- 
phragms into  nine  or  ten  compartments,  but  communicating 


Fig.  155.— Diagram  Illustrative  of  the  Circulation  in  the  Leech. — Dr.  Williams,  a, 
Great  dorsal  vessel ;  c,  ventral  vessel  ;  d,  d,  intercommunicating  vessels  between 
dorsal  and  ventral  trunks ;  e,  e,  lateral  abdominal  trunks  ;  g,  vessels  distributed 
over  the  caecal  appendages  to  the  stomach  ;  /,  /,  /,  loop-shaped  organs  to  the  respira- 
tory vesicles  ;  h,  h,  lateral  branches  to  respiratory  vesicles. 

freely  with  each  other,  and  in  each  of  which  are  two  openings 
communicating  with  the  lateral  caeca. 

Up  to  this  stage  in  development,  then,  in  which  a  very  com- 
plex circulation  is  seen  to  exist,  no  heart  presents,  but  the  ves- 
sels themselves  producing  it,  inclusive,  of  course,  of  the  action 
of  the  polar  forces.  Unless,  forsooth,  the  whole  dorsal  vessel, 
running  the  length  of  the  body,  inclusive  of  the  lateral 
branches,  be  considered  as  such,  which  is  perfectly  absurd — 
the  outcome,  in  fact,  of  the  misleading  and  erroneous  concep- 
tion that  the  animal  circulation  is  based  upon  the  heart,  which 


3SG 


STOMACH   AND    C^ECA   IN   THE  LEECH. 


is  a  means  for  increasing  it  simply,  and  "putting  the  cart  be- 
fore the  horse,"  by  reversing  the  order  in  nature.  Further- 
more, it  would  leave  unexplained  the  action  in  the  other  ves- 
sels ;  consequently,  the  principle  is  \\  rong.  And  as  the  organ  re- 
lates to  the  production  of  force  by  increasing  circulation  in  the 


Fig.  156.— Digestive  Organs  of  the  Leech  (Hirudo  Medicinalis). — Jones,  b,  Pharynx  ;  hr 
i,  interior  of  stomachal  cavity,  exhibiting  the  diaphragms,  with  the  lateral  openings 
in  the  caecal  appendages  (k)  ;  g,  first  pair  of  stomachal  caeca  ;  d,  last  pair,  extending 
backward  on  each  side  of  the  intestine  (e),  which  opens  on  the  dorsal  surface  close  to 
the  terminal  sucker. 

respiratory  organs,  the  absence  of  a  heart  in  the  circulatory  sys- 
tem of  the  annelid es  is  readily  accounted  for  by  the  diffused  res- 
piration which  obtains  here,  for  in  every  part  of  the  circumfer- 
ence of  each  ring  the  blood  is  being  arterialized  as  it  is  being  ren- 
dered venous  ;  hence,  no  heart  is  developed,  the  action  in  the 
dorsal  artery  with  the  lateral  vessels  extending  into  the  capil- 
lary plexuses  of  the  skin,  respiratory-vesicles,  or  branchiae,  as 


NO   HEART   IN   ARENICOLA.  387 

the  case  may  be,  being  sufficient  for  the  purpose  and  subserv- 
ing the  functions  of  a  heart.  But  this  latter  circumstance  is 
made  more  conspicuous  in  arenicola  (Fig.  157),  in  which  the 
circulation  is  simplified.  And  this  beautiful  diagram,  by  the 
distinguished  anatomist  and  naturalist  at  Heidelberg,  will  serve 
to  give  a  distinct  mental  picture  of  the  adaptive  changes  in  the 
vascular  system  to  the  form  of  the  respiratory  apparatus  and 
the  principle  that  obtains  in  the  mechanics  for  increasing 
circulation  in  it  commensurate  with  the  force  which  is  ex- 
pended in  the  activities.  As  will  be  seen,  the  branchial  vessels 
(b,  b)  are  connected  directly  with  the  great  dorsal  and  ventral 
trunks  (d,  v)  the   blood   from   the  latter  passing  into  the 


Fig.  157. — Diagrammatic  Transverse  Section  through  the  Hinder  Half  of  the  Body  of 
Arenicola,  to  show  the  arrangement  of  the  vessels. — Gegenbaur.  D,  dorsal  ;  V, 
ventral  side  ;  n,  ventral  medulla  ;  i,  enteric  cavity  ;  or,  branchiae  ;  v,  ventral  vascu- 
lar trunk  ;  ab,  branchial  vessels  ;  d,  dorsal  vascular  trunk  ;  h,  branch  surrounding 
the  enteric  canal ;  v',  visceral  ventral  vessel. 

branchia  ibr)  by  means  of  the  afferent  vessel  (a),  thence  to 
the  dorsal  artery  through  the  efferent  vessel  (b),  one  upon 
either  side  corresponding  with  the  branchiae  ;  hence,  any  ex- 
pansile action  m  the  branchiae  or  vibratory  motion  tending  to 
increase  circulation  in  the  plexuses  would  necessarily  deter- 
mine the  venous  blood  in  this  direction,  while  the  pumping  ac- 
tion in  the  dorsal  vessel  should  serve  for  aspirating  the  plexuses 
and  propelling  it  through  the  system  ;  since  the  diastoles  pro- 
duce a  suction-force  and  the  systoles  a  driving  force  upon  the 
blood.  And  though  reflux  is  inevitable,  by  reason  of  the  absence 
of  valves,  still  this  is  limited,  from  the  fact  that  the  blood  which 
is  withdrawn  from  the  plexuses  by  the  diastoles  is  instantane- 
ously supplied  by  the  venous  blood  flowing  into  them,  filling 
them  before  the  succeeding  systole  sets  in,  or  nearly  so  ;  so  that 
only  a  small  amount  of  reflux  is  possible.     But  this  mechanics 


388  NO   HEART  IN   ARENICOLA. 

would  not  answer  when  a  heart  comes  in  the  scheme,  for  the 
reason  that  the  force  in  the  latter  would  tend  to  rupture  the 
capillaries  from  the  strain  it  should  occasion  ;  hence  the  dif- 
ferentiation of  valves  for  obviating  reflux.  In  addition  to 
these  vessels,  however,  we  have  now  to  notice  another  arrange- 
ment of  equal  importance,  notably  the  one  relating  to  the 
intestinal  canal,  since  this  is  the  gateway  of  the  nutrient  and 
force-producing  elements.  Here,  for  example,  we  have  two 
lateral  vessels  (h)  that  embrace  the  intestinal  canal  (/)  and  con- 
nect with  the  visceral  ventral  vessel  (vr)  and  the  delicate  plex- 
uses in  the  visceral  parietes  on  the  one  hand,  and  the  dorsal 
vessel  (d)  upon  the  other  ;  and  bearing  in  mind  the  mechanics 
in  the  portal  circulation,  is  it  not  apparent  that  striking  cor- 
respondence subsists  between  them  ;  nay,  that  here  we  have 
indeed  the  very  thing  itself  in  its  nascent  stage  ?  Notably, 
the  visceral  blood  is  collected  by  special  veins  and  delivered 
in  the  systemic  current  (d),  passing  thence  into  the  respiratory 
apparatus  ibr)  by  the  vessels  (bb),  the  pumping  action  in  the 
dorsal  vessel  being  the  analogue  of  that  in  the  heart,  while 
the  contractions  in  the  external  muscular  envelope,  by  in- 
creasing pressure  in  the  viscera,  should  serve  for  expediting 
the  action  ;  not  to  mention  the  suction-force  exerted  by  the 
branchiae  themselves,  which,  of  course,  affects  the  blood 
in  these  vessels  in  similar  manner  as  in  the  corresponding  ves- 
sels to  the  ventral  venous  trunk  (a,  n) ;  and  since  the  animal 
is  in  constant  motion,  the  vermicular  movements  should  in- 
crease the  visceral  circulation  and  respiration  correspondingly. 
It  is  not  fanciful,  since  there  is  definite  arrangement  in  the 
parts,  and  there  is  absolute  necessity  for  such  provision,  in 
order  to  maintain  a  balance  in  the  organism. 

Now,  then,  looking  from  this  diffused  respiration  in  areni- 
cola  to  the  localized  respiration  in  terebella,  where  the 
branchiae  project  from  the  occiput  (Fig.  158,  &,  &),  the  effect 
upon  cardiac  development  is  at  once  made  apparent.  For 
example,  the  great  dorsal  vessel  (m)  is  now  greatly  reduced 
in  size  by  reason  of  the  disappearance  of  the  lateral  branchiae, 
while  the  vascular  oesophageal  collar  (n),  which  receives  all 
the  blood  from  the  intestinal  system,  is  increased  in  size  cor- 
respondingly and  functions  as  an  auricle  to  the  elongated  fusi- 


FIRST   EVIDENCE   OF  A   HEART  IN"   TK  RE  BELL  A. 


:^89 


form  heart  (I)  lying  upon  the  oesophagus  (e)  in  immediate 
relation  with  the  respiratory  organs  (k,  k),  this  portion  of 
the  great  dorsal  artery  taking  on  increased  development  in 
correspondence  with  the  branchiae,  the  other  shrinking  in 
size,  responsive  to  the  changes  in  the  respiratory  organs. 


Fig.  158.— Vascular  System  of  Terebella.— Milne-Edwards.  Showing  the  earliest  indi- 
cations in  cardiac  development  (I),  and  the  relations  it  sustains  to  the  respiratory- 
organs  (k,  k). 

Briefly,  the  haemal  mechanics  is  as  follows :  The  fusi- 
form heart  (d),  which  is  but  slightly  attached  to  the  struct- 
ures on  which  it  rests,  and  suspended,  as  it  were,  in  the 
fluids  of  the  peritoneal  cavity,  sends  off  at  the  anterior  end 
three  lateral  symmetrical  branches  (Fig.  159,  a,  d)  to  the 
branchiae,  the  reduced  continuation  of  the  original  trunk  break- 


390 


PLAN    OF    CIRCULATION    IN   TEREBELLA. 


irig  up  into  minute  vessels  to  the  tentacles,  in  the  hollow  axes 
of  which  each  terminates  in  an  efferent  vessel,  and  which  are 
surrounded  by  the  peritoneal  fluid,  which  penetrates  to  the 
remotest  ends  of  these  exquisite  organs.     But  the  branchiae  are 


Fig.  159. — Plan  of  the  Circulation  in  Terebella. — Dr.  Williams,  a,  Elongated  fusiform 
heart ;  6,  vascular  oesophageal  collar,  which  receives  all  the  blood  from  the  intes- 
tinal system  (auricular)  ;  rf,  the  three  pairs  of  branchial  arteries  to  a  corresponding 
number  of  branchiae  ;  c',  three  pairs  of  branchial  veins  corresponding  with  the 
arterial  conveying  the  aerated  blood  into  the  longitudinal  systemic  trunk  (c,  c)  to  be 
distributed  to  the  intestines  and  body-territories,  three  pairs  of  lateral  branches 
crossing  the  oesophageal  collar  (b)  to  connect  with  the  great  dorsal  vessel ;  /,  frame- 
work of  longitudinal  and  transverse  vessels,  embracing  the  alimentary  canal,  as  in 
all  annelides. 

the  principal  organs  of  respiration,  the  three  pairs  of  large 
lateral  arterial  branches  to  which  terminate  in  a  correspond- 
ing number  of  efferent  vessels  (c' )  that  converge  in  the  great 
ventral  trunk  (<?,  e),  which  extends  to  the  posterior  extremity 
of  the  body,  beneath  the  intestine  and  immediately  above  the 


PL AJST   OF   CIKCULATICXN-   IN    TEllEBhLL.Y.  391 

ventral  chain  of  nervous  ganglia,  but  is  distinct  from,  and 
independent  of,  the  intestinal  system  {/),  and  which  gives  off 
opposite  to  each  ring  a  pair  of  transverse  vessels,  which,  after 
having  supplied  branches  to  the  integument  and  locomotive 
organs  (Fig.  158,  g,  J),  *  bends  upward,  to  be  distributed  over 
the  walls  of  the  intestine,  where  their  ramifications  contribute- 
to  form  the  vascular  network  above  referred  to.  At  the  point 
corresponding  with  the  circular  vessel  (Fig.  159,  b),  the  ventral 
trunk,  however,  gives  off  a  large  branch  to  the  intestine  ;  and 
since  the  blood  flows  out  of  the  branchiae  and  tentacles  through 
this  great  ventral  trunk,  it  follows  that  a  considerable  amount 
of  purely  arterial  blood  must  flow  immediately  into  the  vessels 
of  the  intestine  for  supplying  its  glandular  parietes,  and 
evolving  the  force  which  is  expended  in  the  digestive  func- 
tions. And  here,  again,  we  have  a  distinct  vascular  system, 
the  blood  coursing,  as  in  the  external  envelope,  in  two  direc- 
tions, namely,  longitudinally  and  transversely  or  circularly. 
(In  order  to  understand  the  mechanics,  it  must  be  borne  in 
mind  that  the  intestine  is  surrounded  by  peritoneal  fluid,  the 
organ  being  suspended,  as  it  were,  in  the  peritoneal  cavity  by 
means  of  limited  bands  proceeding  from  it  to  the  external 
envelope,  tying  the  two  cylinders  together,  at  the  same  time, 
however,  permitting  them  to  move  freely,  the  one  within  the 
other.) 

Finally,  for  re-collecting  the  blood  in  these  independent 
tubes,  we  have  the  vessels  communicating  with  the  great 
circular  auricle  in  immediate  relation  with  the  fusiform 
heart  (Fig.  159,  b,  a),  and  by  means  of  the  diastoles  and  sys- 
toles occurring  in  these  vessels,  the  blood  is  pumped  into  the 
respiratory  apparatus,  thus  completing  the  circle. 

But  the  influence  exerted  upon  cardiac  development 
from  localizing  respiration  has  more  forcible  illustration 
in  decapods  (Fig.  160).  Here  the  venous  blood  is  col- 
lected from  body  territories  and  locomotive  organs  in  large 
venous  sinuses  at  the  bases  of  the  branchiae  (a,  a)  thence  by  a 
special  vessel  (e,  e)  analagous  to  the  pulmonary  artery  to 
the  branchiae    (one    for   each   gill),    to    be    distributed  over 

*  The  intestine  is  pulled  aside  in  order  to  exhibit  these  vessels. 


392       SHOWING   POSITION   OF   THE   HEART   IN   DECAPODS. 

the  innumerable  minute  laminse  of  the  gills,  whence  it  is  re- 
collected by  the  branchial  veins  (/,  /'),  and  transmitted  to  the 
heart  (g).  The  branchial  arteries  follow  the  outer  margin,  and 
the  returning  veins  the  inner  margin  of  the  gills,  and  the  united 
trunks  of  the  latter  vessels  convey  the  arterialized  blood,  by  a 
single  orifice  on  each  side,  into  the  large  median  muscular 
ventricle  (g).  As  in  other  articulata,  it  is  situate  in  the 
middle  of  the  back,  as  seen  in  the  lobster  (Fig.  161,  c), 
and  consists  of  a  single  systemic  muscular  cavity,  most  con- 
centrated in  form  in  the  decapods,  and  generally  elongated  on 
the  inferior  orders ;  its  thick  parietes  are  composed  of  inter- 


Fig.  160.— Showing  Relation  of  the  Heart  to  the  Gills  in  Decapods,  Maja  Squinado. — 
Grant,  a,  a,  Capacious  lateral  sinuses  at  the  bases  of  the  branchiae  ;  b,  inferior 
abdominal  veins  ;  c,  superior  abdominal  veins  ;  e,  e,  branchial  arteries  originating 
from  these  wide  sinuses  :  /,  /,  branchial  veins  ;  g,  the  large  median  muscular  ventri- 
cle ;  i,  i,  h,  anterior  arterial  branches  ;  k,  the  great  posterior  median  systemic 
artery. 

laced  muscular  fibres,  they  present  internal  fleshy  columns, 
there  are  semilunar  valves  at  the  orifices  of  the  great  vessels, 
and  the  heart  is  connected,  as  usual,  with  the  neighboring 
parts  by  muscular  bands.  From  the  anterior  and  upper  mar- 
gin of  the  heart  arise  three  arterial  trunks  (/,  /,  h),  the  two 
lateral  of  which  send  branches  to  the  genital  organs  and  the 
stomach,  and  terminate  in  the  two  antennal  arteries,  proceed- 
ing to  the  outer  and  inner  pair  of  these  organs,  and  the  median 
vessel  (h),  advancing  over  the  stomach  to  the  pedunculated 
eyes,  divides  into  the  two  ophthalmic  arteries,  which  supply 
these  organs.  From  the  lower  and  anterior  part  of  the  heart,  the 
hepatic  arteries  originate  by  a  single  or  double  trunk,  accord- 
ing to  the  divided  condition  of  the  liver,  through  which  they 


SHOWING   POSITION"   OF   THE  HEART  IN   LOBSTER. 


393 


ramify ;  and  from  the  lower  and  posterior  part  of  the  same 
muscular  cavity  arises  the  great  sternal  artery,  which,  after 


Fig.  161. — Diagram  of  the  Circulatory  System  of  a  Lobster.  — Gegenbaur.  0,  eye  ;  ae, 
lateral  antennas  ;  br,  branchiae  ;  c,  heart ;  pc,  pericardium  ;  ao,  median  anterior 
aorta  ;  aa,  hepatic  artery  ;  op,  posterior  artery  of  the  body  ;  a,  trunk  of  the  ventral 
artery  ;  v,  ventral  venous  sinus  ;  v  br,  branchial  veins.  The  arrows  show  direction 
of  the  blood  current. 

descending  to  the  sternum  and  dividing  into  an  anterior  and 
posterior  trunk,  supplies  most  of  the  musculo-cutaneous  parts 


394  PLAN   OF   THE   CIRCULATION. 

on  the  ventral  region  of  the  body,  and  gives  off  laterally  the 
brachial  arteries  to  the  locomotive  organs.  From  the  middle 
of  the  posterior  margin  of  the  heart  is  given  off  the  great  pos- 
terior median  systemic  artery  (k),  which,  extending  backward 
along  the  median  and  dorsal  part  of  the  trunk,  sends  off  nu- 
merous branches  on  each  side  to  the  neighboring  organs,  and 
bifurcates  over  the  colon  before  distributing  its  terminal 
branches  on  the  muscles  of  the  tail.  The  venous  blood  of  the 
system  is  returned  to  the  great  ventral  and  branchial  sinuses 
and  transmitted  through  the  gills,  and  when  arterialized  is  con- 
veyed upward  to  the  dorsal  part  of  the  trunk  by  the  branchial 
veins,  to  be  poured  into  the  cavity  of  the  heart  by  two  orifices 
on  its  upper  surface.  The  venous  and  arterial  openings  of  the 
heart  are  provided  with  valves  (Grant).  Thus,  we  have  cardiac 
development  coming  into  prominence  in  connection  with  the 
differentiation  of  a  local  respiratory  apparatus,  and  growing 
more  and  more  perfect  with  progress  in  development.  But  the 
point  to  which  attention  is  specially  directed  is  the  location  of 
the  heart  (g)  itself,  which  is  between  the  gills  and  upon  the 
distal  side,  so  to  speak,  of  this  circulation,  while  the  large 
venous  sinuses  (a,  a),  from  which  the  pulmonary  arteries  (e,  e) 
take  their  origin,  are  upon  the  opposite  side  ;  the  one  serving 
for  aspirating  and  propelling  the  venous  blood  in  the  organs, 
the  other  for  aspirating  it  thence  and  propelling  it  through  the 
systemic  vessels.  Here,  then,  is  clearly  indicated  the  mechani- 
cal principle  in  the  perfect  heart,  the  right  side  serving  for 
aspirating  the  blood  in  the  veins,  and  compelling  it  in  the 
alveolar  capillaries,  the  left  aspirating  the  capillaries  and  com- 
pelling the  blood  in  the  systemic  vessels,  only  that  one  must 
not  lose  sight  of  the  action  in  the  lungs  themselves,  the  same 
remark  applying  for  every  stage  in  development. 

In  the  lobster  (Fig.  161),  the  branchiae  (br)  are  increased  to 
six  or  more  pairs  ;  accordingly,  we  have  a  similar  increase  in 
the  number  of  pulmonary  veins  (v  br),  conveying  the  blood 
to  the  heart  (c),  which  occupies  the  dorsal  region,  as  before 
remarked  ;  while  the  great  ventral  sinus  (v)  at  the  bases  of  the 
branchiae  sends  out  a  corresponding  number  of  pulmonary 
arteries.  But  we  have  now  to  notice  an  advance  in  car- 
diac development,  notably  the  differentiation  of  a  pericardial 


PLAN   OF   THE   CIRCULATION".  395 

sinus  (pc),  in  which  the  pulmonary  veins  terminate,  and  from 
which  the  blood  flows  into  the  cardiac  chamber  through 
three  pairs  of  cleft-like  openings,  symmetrically  arranged  and 
guarded  by  valves,  which  project  into  the  heart,  for  obviating 
reflux  during  systole  ;  but  in  the  fishes  (in  which  a  true  auricle 
is  formed)  is  converted  into  the  pericardium,  lubricating  the 
organ  and  facilitating  the  rhythmical  expansions  and  con- 
tractions, in  consequence  making  it  more  efficient.  In  other 
respects,  the  circulation  is  fundamentally  the  same  as  in 
maja  squinado  (Fig.  160).  And  looking  from  this  to  the 
corresponding  stages  of  development  in  the  tracheata,  we  find 
similar  arrangements  obtain  in  them  also.  Thus,  in  arach- 
nida,  for  example,  the  respiratory  apparatus  is  in  the  abdo- 
men, the  stigmata  opening  npon  the  sides  and  front  of  the 
a,bdomen,  while  the  pulmonary  sacs,  which  correspond  with 
the  anterior  respiratory  orifices,  are  in  relation  with  the  larg- 
est division  of  the  heart,  which  is  located  in  the  dorsal  re- 
gion and  divided  into  three  compartments,  each  one  of  which 
is  furnished  with  a  pair  of  cleft-like  openings,  symmetrically 
arranged  on  the  dorsal  surface  and  guarded  by  a  pair  of 
valves.  The  arterialized  blood  passes  into  the  cardiac  chambers 
through  these  clefts,  thence  through  the  systemic  and  lateral 
branches ;  the  great  anterior  artery  passing  through  the 
cephalothorax  to  supply  the  anterior  portions,  and  the 
smaller  posterior  vessel  to  the  viscera  and  walls  of  the  ab- 
domen. But  the  tracheae  are  distributed  like  blood-vessels 
through  the  body,  bringing  every  portion  in  immediate  rela- 
tion with  the  air,  which  is  pumped  in  and  out  of  the  canals 
for  effecting  interchange  of  the  gases,  each  stigma  guarded 
by  a  valve,  which  is  opened  and  closed  for  the  purpose  by 
means  of  special  muscles  ;  while  the  blood  passes  to  and  from 
the  respiratory  organs  by  means  of  lacunar  passages,  reaching 
the  heart  in  this  manner  and  passing  through  the  ostia  during 
the  diastoles,  drawn  by  the  suction-force  in  the  heart.  The 
pericardial  sinns  is  also  absent ;  but  not  in  scorpionea  (which 
correspond  in  development  with  the  lobster),  in  which  the 
heart  is  mnch  larger  than  in  arachnida,  in  correlation  with 
the  form  of  the  body  and  the  number  of  the  tracheae,  and 
surrounded  by  a  pericardial   sinus ;   is    divided    into    eight 


396  PLAN   OF   THE   CIRCULATION. 

chambers,  with  a  pair  of  dorsal  clefts  leading  into  each  cham- 
ber, and  guarded  by  valves  which  project  into  the  cavity. 
"  Arterial  vessels  are  given  off  from  the  anterior  as  well  as 
from  the  posterior  end  of  the  heart,  of  which  they  are  direct 
prolongations  ;  the  anterior  one,  the  aorta,  enters  the  cephalo- 
thorax,  while  the  hinder  one  runs  to  the  tail.  In  addition  to 
these,  a  number  of  lateral  arteries  are  given  off  close  to  the  ve- 
nous ostia,  and  are  distributed  to  the  neighboring  organs. 
Two  of  the  numerous  branches  given  off  by  the  aorta  form  a 
vascular  ring  around  the  oesophagus,  where  an  artery  runs 
back  (arteria  supraspinalis)  on  and  as  far  as  the  end  of  the 
ventral  nerve-chord  ;  this  artery  gives  off  a  large  number  of 
branches.  The  venous  blood  is  collected  in  a  receptacle  which 
lies  directly  on  the  ventral  surface,  just  as  in  the  higher  Crus- 
tacea ;  from  this  it  is  carried  to  the  respiratory  organs.  Be- 
fore the  blood  from  them  gets  to  the  heart,  it  passes  into  the 
pericardial  sinus  "  (Gegenbaur)  Italics  are  added.  It  is  thus 
made  manifest  that  the  vascular  system  is  set  to  the  respira- 
tory apparatus,  in  which  it  all  converges  as  the  spokes  in  the 
u  hub"  of  a  wheel,  being  the  basis,  so  to  speak,  of  the  organ- 
ism ;  while  the  heart  shows  adaptive  changes  in  correspond- 
ence with  the  local  mechanics. 

In  conchifera  are  two  pairs  of  large  laminated  branchiae, 
while  the  heart,  which  is  still  more  developed,  is  at  the  base 
of  these  (Fig.  16J,  n,  o),  in  a  cavity  between  the  great  adduc- 
tor muscle  (I,  m)  and  the  folds  of  intestine.  It  possesses  a 
true  pericardium,  and  consists,  in  the  species  under  considera- 
tion, of  two  distinct  chambers — an  auricle  and  ventricle.  The 
auricle  (163,  b)  is  large,  and  the  walls  extremely  thin,  the 
delicate  muscular  fasciculi  and  transparent  membranes  per- 
mitting the  blood  to  be  seen  through  them.  It  receives  the 
blood  from  th°s  branchiae  by  two  large  venous  trunks,  in  which 
the  branchial  veins  converge  (e,  g,  7i),  and  transmits  it  to  the 
ventricle  (d)  through  two  intermediate  canals  (c),  whence  it  is 
propelled  through  the  body  by  the  arterial  vessels  {n,  o,  p), 
a  vessel  of  considerable  size,  passing  at  once  in  the  ad- 
ductor muscle  for  producing  the  force  which  is  expended  in 
operating  the  valves.  The  veins  pass  directly  into  the 
branchiae  (m,  n,  i,f)  without  the  intervention  of  a  sinus,  the 


POSITION   OF  THE   HEAET   IN  THE   OYSTEE. 


397 


Fig.  162. — Alimentary  Canal  of  the  Oyster  (Ostrea  edulis). — Jones,  a,  The  stomach 
laid  open  ;  d,  the  liver  ;  b,  c,  d,  f,  convolutions  of  the  intestine  ;  gr,  anal  aperture  ; 
n,  o,  auricle  and  ventricle  of  the  heart ;  I,  m,  adductor  muscle  ;  h,  k,  lobes  of  the 
mantle,  divided  to  show  the  large  venous  canals  at  the  base  of  the  branchiae. 


Fig.  163. — Heart  and  Respiratory  System  of  the  Oyster. — Jones,  a,  Portion  of  the 
mantle ;  b,  auricle,  and  d,  ventricle  of  the  heart ;  m,  n,  i,  f,  veins  bringing  the 
blood  from  the  body  to  the  branchiae  ;  e,  gr,  h,  principal  trunks  returning  the  blood 
from  the  branchiae  to  the  heart  ;  o,  p,  J,  aorta,  giving  off  arteries  to  the  viscera. 


398  INCREASE    IN    CARDIAC    DEVELOPMENT. 

action  in  the  branchiae  appearing  to  be  sufficient  for  the  pur- 
pose, while  the  heart  aspirates  the  oxygenated  blood  and 
pumps  it  in  the  tissues,  the  adductor  muscle  getting  a  large 
share.  While  the  above  description  will  hold  good  in  all 
essential  points  for  every  family  of  conchiferous  Mollusca,  yet 
important  modifications  in  the  structure  of  the  heart  and 
arrangement  of  the  blood-vessels  are  met  with  in  different 
genera  ;  most  generally  in  consequence  of  the  increase  in  size, 
since  this  requires  more  force  for  operating  the  valves,  with  a 
corresponding  increase  in  the  respiratory  surface  and  in  cardiac 
structure.  Accordingly,  we  have  two  auricular  cavities,  one 
for  each  pair  of  branchial  lamellae,  placed  symmetrically  on 
the  two  sides  of  an  elongated  fusiform  ventricle,  into  which 
both  auricles  empty  themselves,  while  in  the  forms  remarkable 
for  their  breadth,  e.  g.,  area,  there  are  not  only  two  auricles, 
but  two  ventricles  likewise,  placed  at  the  base  of  each  pair  of 
gills,  upon  opposite  sides  of  the  body,  each  receiving  the 
blood  from  the  branchiae  to  which  it  belongs,  and  propelling 
it,  through  vessels  common  to  both  hearts,  to  all  parts  of  the 
system ;  and  showing  conclusively  the  need  of  arterialized 
blood  and  that  the  form  of  the  respiratory  apparatus  con- 
ditions cardiac  development. 

In  the  fishes  the  heart  is  carried  forward  to  the  base  of  the 
skull  (Fig.  164,  a),  for  the  reason  that  the  oxygenating  appa- 
ratus is  also  here,  the  branchiae  being  articulated  with  the 
base  of  the  skull.  Hence  this  phenomenon.  And  the  prin- 
ciple in  animal  mechanics  borne  in  mind,  that  fact  explains 
itself. 

The  form  of  the  heart  varies  in  the  fishes,  but  may  be  briefly 
described  as  consisting  of  an  auricle  and  ventricle  developed 
in  the  venous  system  in  immediate  relation  with  the  branchial 
(Fig.  165,  a,  v,  bf),  by  means  of  which  the  venous  blood  is 
pumped  in  the  branchiae,  whence  the  arterialized  blood  flows 
through  the  aorta  {ad)  to  the  body-territories  ;  the  early  fore- 
shadowing of  which  is  seen  in  terebella  (Fig.  158),  in  which 
the  tubular  auricle  (n)  and  fusiform  heart  (7),  with  the  great 
efferent  vessel  (Fig.  159,  c,  c),  conveying  the  arterialized  blood 
from  the  branchiae  to  the  body-territories,  are  archetypal.  A 
momentary  glance  into  the  mechanics  may  not  come  amiss. 


POSITION   OF  HEART  IIST   THE  FISHES. 


399 


Fig.  164. — Viscera  of  the  Shark,  in  situ,  a,  The  heart ;  b,  gill-openings  ;  c,  c,  c,  lobes 
of  the  liver  ;  d,  e,  f,  h,  alimentary  canal  ;  i,  appendage  to  the  intestine  ;  g,  biliary 
duct;  n,  the  testis  ;  o, p,  vas  deferens  ;  k,  intromittent  organ;  s,  openings  com- 
municating with  the  peritoneal  cavity  ;  I,  claspers. — Owen. 


400  PLAN   OF  THE   CIRCULATION   IN   FISHES.. 

For  force  and  speed  of  locomotion  (the  one  involving  the 
other),  the  fishes  eclipse  all  the  branchiata  ;  but  since  thia  in- 
volves the  more  rapid  oxygenation  of  the  blood,  the  special 
arrangements  which  obtain  for  effecting  it,  are  at  once  made 
intelligible.  For  this  purpose,  the  respiratory  surface  is  am- 
plified by  the  formation  of  branchial  leaflets  projected  from  the 
gill  arches  (Fig.  16G),  while  the  branchial  arteries  rapidly  divide 
and  subdivide  nntil  they  resolve  themselves  into  microscopic 
capillaries,  pervading  every  portion  of  the  leaflets,  and  covered 
only  by  a  thin  tessellated  epithelium,  so  that  free  interchange 
of  the  gases  is  readily  effected  between  the  blood  and  the 
water,  which  is  pumped  over  them  by  the  action  in  the  mouth 
and  opercula,  and  which  is  increased  and  diminished  in  corre- 


Fig.  165. — Head  of  an  Embryonic  Teleostean,  with  the  rudiments  of  the  vascular  system 
(diagrammatic). — Gegenbaur.  a,  Auricle  ;  v,  ventricle  ;  abr,  branchial  artery  ;  c, 
carotid  ;  ad,  aorta  ;  s,  branchial  clefts  ;  n,  nasal  pit  ;  sv,  sinus  venosus  ;  dc,  ductus 
Cuvieri. 

spondence  with  the  physiological  requirements.  Now,  then, 
the  point  to  which  attention  is  specially  directed  concerns  the 
mechanical  principle  which  applies  to  this  pumping  action  in 
the  mouth  and  opercula,  and  the  effect  it  has  upon  circulation 
in  the  fishes,  the  venous  especially.  Thus,  pressure  being  uni- 
form, it  is  obvious  that  with  each  expansion  in  the  mouth  and 
opercula  for  reducing  pressure  in  the  branchiae,  should  produce 
afflux  of  the  venous  blood  in  the  organs  simultaneously  with 
the  afflux  of  fresh  water,  since  pressure  is  transmitted  through 
and  through  the  body  upon  all  the  organs  the  same  as  in  air- 
breathers,  the  fluids  flowing  from  high  to  low  pressure  in  con- 
formity with  universal  law  ;  while  during  contraction  for  com- 
pelling the  respired  water  out  of  the  organs  by  increasing 
pressure,  that  this  should  also  tend  to  drive  the  oxygenated 
blood  into  the  aorta,  since  the  valves  in  the  pulmonary  artery 


RESPIRATORY   NERVES   IN   FISHES. 


401 


would  obviate  reflux,  the  effect  being  the  same  as  in  air- 
breathers,  only  there  is  no  auricle  or  ventricle  in  the  left  side 
for  assisting  the  action. 

For  evolving  the  force  which  is  expended  in  the  activities  the 
grand  object,  of  course,  is  to  bring  the  venous  blood  into  the 
branchiae  in  correspondence  with  the  afflux  of  fresh  water,  and, 
reasoning  from  analogy,  can  it  be  doubted  for  a  single  moment 
but  that  the  heart  is  coordinated  with  the  branchiae  the  same 


Pig.  166. — Brain  and  Cerebral  Nerves  of  the  Perch  (after  Cuvier).  a,  The  cerebellum  ; 
6,  cerebrum  ;  c,  olfactory  ganglia  ;  i,  bulbous  commencement  of  the  olfactory  nerve; 
o,  o,  olfactory  nerve,  terminating  in  the  nasal  capsule';  n,  optic  nerve  ;  p,  q,  third, 
fourth,  and  six  pairs  of  nerves,  appropriated,  as  in  Man,  to  the  muscles  of  the  eye- 
ball ;  a,  ophthalmic  branch  of  the  fifth  pair  ;  (3,  superior  maxillary  branch  of  ditto ; 
0,  inferior  maxillary  branch  of  ditto  ;  ju,  opercular  branch  ;  |,  branch  of  the  fifth 
pah-,  mounting  upwards  to  join  @,  a  branch  from  the  eighth  pair,  running  to  supply 
the  dorsal  region  of  the  body  ;  s,  s,  auditory  nerve  ;  t,  t',  nerves  belongmg  to  the 
eighth  pair  ;  iv,  z,  nerves  answering  to  the  spinal  recurrent. 

as  in  the  lungs  of  the  air-breather,  the  principle  being  the 
same.  But  for  correct  appreciation  of  the  mechanics  it  will  be 
necessary  to  bring  out  more  clearly  the  action  of  another  force 
which  applies  here,  and  forming,  as  it  were,  the  foundation  of 
the  mechanism ;  namely,  the  density  of  the  medium,  which 
tends  to  equilibrate  gravitation,  at  the  same  time  that  it  in- 
creases pressure. 

For  example,  when  water  is  poured  out  of  a  vessel  in  the 
atmosphere,  it  rushes  with  great  impetuosity  to  the  ground 


402  RESPIRATORY   NERVES   IN   FISHES. 

from  the  action  of  gravitation,  the  atmosphere  offering  but 
little  obstruction  by  reason  of  the  rarity  of  the  medium  ;  but 
when  poured  in  the  ocean  it  is  upheld  in  the  surface  waters, 
and  the  blood  being  but  slightly  heavier  than  water,  it  follows, 
that  in  the  case  of  the  fishes  the  blood  needs  only  to  be  guided 
to  and  from  the  branchiae  by  the  vessels  in  order  to  effect 
oxygenation,  while  the  pumping  action  in  the  head  by  reason 
of  the  high  pressure  in  the  body  should  compel  afflux  and 
efflux  of  the  blood  in  the  branchiae.  Hence,  the  undeveloped 
condition  of  the  vessels  in  the  fishes,  together  with  the  fact 
that  circulation  in  the  tissues  is  carried  on  through  canalicular 
spaces  under  the  play  of  the  polar  forces,  vis  a  tergo,  by  reason 
of  the  absence  of  a  ventricle  in  the  arterial  system,  being  re- 
duced to  a  minimum.  The  grand  object,  as  before  remarked, 
is  to  arterialize  the  blood,  which  increases  polar  force  ;  and  for 
effecting  this  commensurate  with  the  physiological  require- 
ments we  have  not  only  the  great  pumping  action  in  the  head, 
but  a  more  extensive  movement  (reasoning  from  analogy)  is 
also  taking  place — namely,  a  pumping  movement  in  the  abdo- 
men synchronous  with  respiration  for  increasing  the  portal 
circulation,  in  order  to  maintain  correspondence  between  the 
absorptive  processes  in  the  canal  and  respiration ;  the  same 
principle  applying  as  in  land  animals,  otherwise  a  balance 
could  not  be  maintained. 

Then,  again,  judging  from  the  celerity  with  which  the  air- 
bladders  are  expanded  and  contracted,  which  would  involve 
corresponding  movements  in  the  walls  of  the  abdomen  for 
effecting  these  actions,  we  may  very  readily  understand  how 
this  increase  in  the  portal  circulation  could  be  produced. 

Last,  but  not  least,  the  correlation  of  the  nerves  in  the 
respiratory  centre  (Fig.  166,  g),  extending  thence  over  the 
sides  of  the  abdomen  (Fig.  167,  2,  3,  4),  plainly  indicate  how 
the  parts  are  coordinated  with  respiration.  Kay,  not  the  ab- 
domen only,  but  the  tissues  of  the  entire  body  as  well,  since 
intimate  nervous  connections  bind  all  the  parts  to  respiration ; 
the  fins  (V,  P,  A),  tail  and  all  the  muscles  in  the  body  being 
intimately  connected  with  the  medulla  oblongata  and  respira- 
tory centre  by  means  of  the  "nervus  lateralis"  (1,  2,  3,  4),  where- 
by coirespondence  is  readily  produced,  and  respiratory  move- 


RESPIKATOEY  NERVES   IN"  FISHES.  403 

ment  made  to  pulsate  through  and  through  the  body.  Thus, 
by  means  of  the  nerves,  the  whole  is  unified  with  the  action  in 
the  branchiae,  which  should  be  so,  for  the  very  obvious  rea- 
son that  it  all  relates  to  the  evolution  of  force,  for  which  respi- 
ration is  the  relative  adjustment,  compelling  the  commerce  in 
the  organism  in  the  measure  of  the  physiological  requirements. 
The  very  intimate  connection  subsisting  between  respiration 
and  the  voluntary  movements  is  forcibly  exemplified  in  the 
nervous  apparatus  in  fishes.  For  example,  the  eighth  pair 
undergoes  development  in  proportion  to  the  size  and  extent  of 
the  branchial  surface,  at  the  same  time  sending  branches  to 
the  viscera;  while  externally  it  projects  the  great  lateral 
branches  to  the  dorsal  and  ventral  regions  (Fig.  167, 1,  2,  3,  4), 
which  connect  with  the  spinal  nerves  by  means  of  the  dorsal 
and  ventral  roots  (Fig.  16b,  a,  v).  In  other  words,  the  dorsal 
root  of  the  spinal  nerves  sends  a  filament  upward  (a),  which 
joins  a  ventral  filament  (b)  from  the  preceding  nerve,  forming 
the  "ramus  dorsalis,"  and  sends  two  filaments  downward  (c), 
which  unite  together  to  form  the  "ramus  ventralis,"  each 
of  which  joins  the  eighth  pair,  thus  connecting  and  unifying 
the  actions  throughout  with  respiration.  This,  together  with 
the  fifth  pair  (connected  with  mouth  and  opercula),  send  out 
similar  branches  to  the  muscular  envelope. 

How  otherwise  explain  these  comprehensive  nervous  combi- 
nations and  the  great  development  of  the  pneumogastric 
nerves  ? 

The  following,  from  the  painstaking,  indefatigable  and 
venerable  Owen  (now  in  the  nineties),  will  serve  to  show  how 
intimately  every  portion  of  the  body  in  the  fishes  is  connected 
with  the  respiratory  centre : 

"The  vagus  (Fig.  166,  tf)  has  a  development  proportional  to 
the  extent  and  complexity  of  the  branchial  or  respiratory  appa- 
ratus, and  is  usually  larger  than  the  trigeminal ;  it  rises  from 
the  restiform  tract  forming  the  side  of  the  medulla  oblon- 
gata, and  commonly  from  a  specially  developed  lobe,  and 
is  distributed  to  the  branchial  apparatus,  the  pharynx  and 
pharyngeal  arches,  the  oesophagus  and  stomach ;  it  sends 
filaments  to  the  heart,  and  to  the  air-bladder  when  this  exists, 
and  it  forms,  or  helps  to  form,  the  'nervus  lateralis'   (Fig. 


404  INSPIRATORY    NERVEs    IX    FISHES. 

167).  A  branch  of  the  vagus  ascends  forward  to  join  the  fifth 
in  forming  the  dorsal  division  (2)  of  the  'nervus  lateralis,' 
which  escapes  by  a  foramen  in  the  parietal  bone  ;  the  rest  of 
the  fifth  emerges  from  the  skull  by  a  hole  (carp)  or  a  notch 
(cod)  of  the  alisphenoid.  The  lateral  nerve  in  the  cod  is 
formed  chiefly  by  the  fifth,  and  receives  only  a  slender  fila- 
ment of  the  vagus.  In  the  carp,  the  vagus  chiefly  forms  the 
lateral  nerve.  In  the  cod,  the  lateral  nerve  first  sends  off  a 
branch  (1),  which  runs  along  the  sides  of  the  interneural 
spines,  receiving  branches  from  all  the  spinal  nerves  ;  it  then 
curves  down  along  the  scapular  arch,  gives  branches  to  the 
pectoral  (p)  and  ventral  (v)  fins,  supplies  the  great  lateral 
muscular  masses  (2),  and  the  mucous  canal  (-5),  and  sends  a 
nerve  (4),  to  the  interhsemal  spines,  which  communicates  with 
filaments  from  the  corresponding  spinal  nerves ;  both  inter- 
neural and  interhsemal  branches  terminate  in  the  plexus  sup- 
plying the  caudal  fin  ;  thus  all  the  locomotive  members  are 
associated  in  action  by  means  of  the  nervi  laterales.  .  .  . 
In  the  carp  and  herring,  the  vagal  '  ramus  lateralis'  sends  off 
a  strong  branch  to  the  dorsal  fin ;  in  the  garpike,  it  sends,  as 
in  the  cod,  branches  to  the  pectoral  and  ventral  fins  ;  it  dis- 
tributes other  branches  to  the  skin  and  mucous  ducts  ;  and 
some  of  these,  in  most  fishes,  anastomose  with  branches  of  the 
spinal  nerves  (Fig.  167).*  In  the  perch,  there  are  two  'nervi 
laterales,'  above  described,  and  the  proper  lateral  nerve ; 
this  is  formed  exclusively  in  the  vagus,  and  divides  into 
a  superficial  branch,  supplying  the  lateral  line,  and  a  deep- 
seated  branch,  communicating  with  the  spinal  nerves,  and 
supplying  the  myocommatal  aponeuroses  and  the  skin.  .  .  . 
'1  he  vagus  sends  branches  to  the  head  and  opercula  branches  to 
the  gill-covers.  The  usually  double  roots  of  the  nervus  vagus 
pass  out,  in  most  fishes,  by  a  single  foramen  in  the  occipital 
bone.  The  fore  part  of  the  root  is  the  largest,  and  is  gang- 
lionic ;  it  is  the  true  pneumogas trie  nerve,  supplying  the  gills, 
pharynx,  heart  and  stomach,   and  sending  filaments  to  the 

*  Comparative  Anatomy  and  Physiology  of  Vertebrates,  Vol.  I.,  Fishes  and 
Reptiles,  pp.  303-309,  by  Richard  Owen,  F.  R.  S..  Superintendent  of  the  Natural 
History  Department  of  the  British  Museum ;  Foreisn  Associate  of  the  Insti- 
tute of  France,  etc. 


RESPIRATORY   MOVEMENT   UNIVERSAL   IN    FISHES.         405 

septum  dividing  the  branchial  from  the  abdominal  cav- 
ity. .  .  .  Each  vagal  nerve  of  the  sturgeon  equals  the 
spinal  cord  in  size,  and  rises  by  numerous  roots. 

"  The  peculiar  combination  of  the  dorsal  and  ventral  roots  of 
the  spinal  nerves  in  osseous  fishes  is  well  seen  in  the  cod.  The 
dorsal  root  sends  a  filament  (Fig.  168,  a)  upward,  which  joins 
a  ventral  filament  (b),  from  the  preceding  nerve,  and  forms  the 
ramus  dorsalis  (d)  ;  the  dorsal  root  sends  two  filaments  (c) 
downward,  which  unite  together,  and  with  a  ventral  filament 
(e)  of  the  same  nerve  to  form  the  'ramus  ventralis'  (v).  The 
filament  of  the  ventral  root  sent  to  the  ramus  dorsalis  of  the 
succeeding  nerve  perforates  the  lower  division  of  the  dorsal 
root  of  its  own  nerve.  Thus,  each  spinal  nerve  forms  a 
'  ramus  dorsalis '  (Fig.  167,  10),  and  a  '  ramus  ventralis ' 
(8);  the  ramus  dorsalis  includes  a  sensory  filament  of  its  own 
nerve,  and  a  motor  filament  of  the  antecedent  nerve  ;  the 
'  ramus  ventralis '  is  formed  by  a  motory  and  a  sensory  fila- 
ment of  its  own  nerve ;  both  rami  '  ventrales '  and  '  dorsales ' 
are  associated  together,  and  with  the  vagal  and  trigeminal 
nerves  through  the  medium  of  the  great  'nervus  lateralis'  " 
(Figs.  167,  1,  8).  It  will  thus  be  seen  that  the  entire  body  of 
the  fish  is  intimately  connected  with  the  medulla  oblongata 
and  respiratory  centre  for  unifying  the  action  with  respiration, 
the  branchial  apparatus,  heart  and  viscera  being  alike  in- 
cluded, to  the  end,  that  force  may  be  evolved  in  the  measure  of 
the  physiological  requirements,  for  maintaining  a  balance  in 
the  organism  ;  otherwise  impossible. 

Finally,  and  looking  from  these  e^ensive  connections  sub- 
sisting between  the  respiratory  centre  and  the  body-tissues 
for  compelling  correspondence  to  the  little  force  pump  in  the 
venous  system  at  the  base  of  the  skull  in  immediate  relation 
with  the  branchise  (Fig.  164,  a),  can  it  be  doubted  for  a  single 
moment  but  that  the  entire  mechanics  is  based  upon  respira- 
tion for  pumping  the  commerce  in  the  organism  commen- 
surate with  the  physiological  requirements,  while  the  heart 
assists  in  the  action  by  increasing  circulation  of  the  blood  in 
the  branchiae  for  bringing  this  in  correspondence  with  the 
water  flowing  through  the  organs,  produced  by  the  great 
pumping  action  in  the  mouth  and  opercula  but  involving  the 


406 


RESPIRATORY    MOVEMkNT    UNIVERSAL   IN    FISIIES. 


Fig.  167. — Nerves  to  the  Cod  (Gadus  Morrhua)  ;  reduced. — Illustrations  of  the  Com- 
parative Anatomy  of  the  Nervous  System,  4to.  1835.  By  Joseph  Swan.  1, 2,  3, 
4,  "  Nervi  laterales,"  formed  by  the  union  of  the  fifth  and  eighth  pairs,  showing  how 
they  connect  with  the  "rami  ventrales "  and  " rami  dorsales "  of  the  spinal  nerves, 
so  as  to  unify  the  parts  with  the  medulla  oblongata  and  respiratory  centre  ;  v, 
ventral  fin,  showing  the  nerves  (5,  6)  running  into  it  from  the  u  nervus  lateralis  ;'* 
P,  pectoral  fin,  similarly  supplied  ;  A,  tail  fin,  showing  source  of  nervous  supply  ; 
10,  "  ramus  dorsalis  ;"  8,  "ramus  ventralis  ;"  9,  roots  of  ramus  dorsalis  and  ramus 
ventralis,  the  whole  connecting  with  the  respiratory  centre. 


Fig.  168.— Connection  of  Spinal  and  Lateral  Nerves,  Cod.—  Tb.     a,  b,  Roots  of  ramus 
dorsalis  ;  d,  ramus  dorsalis  ;  •»,  "  ramus  ventralis  ;"  c,  c,  roots  of  ramus  ventralis. 


THE  EEL  AMPHIBIOUS.  407 

entire  organism  %  Taking  all  this  into  consideration,  then, 
together  with  the  innumerable  local  actions  which  are  in- 
creased and  diminished  upon  occasion,  the  statement  that  "the 
heart  is  the  force  in  the  circulation  is  too  absurd  to  be  seriously 
entertained."  NicM  zu  glauben.  The  heart  has  enough  to 
do  filling  the  role  in  its  functions,  and  there  is  no  occasion  for 
despoiling  the  vast  autonomy  in  the  tissues  in  order  to  make  it 
appear  marvelous,  for  it  is  not  more  so  than  the  other  organs, 
each  doing  its  own  special  work  in  the  vital  drama  for  which 
it  alone  is  capable,  while  all  of  life  is  marvelous  and  passing 
strange,  a  mystery  of  mysteries. 

In  the  eel,  the  branchiae  are  of  loose  texture  and  very  vas- 
cular, bleeding  freely  when  even  gently  rubbed  between  the 
fingers,  the  capillaries  which  cover  the  surface  in  a  dense 
plexus  breaking  readily  under  the  friction ;  whereas,  in 
fishes,  the  organs  are  of  dense  texture  and  can  endure  an 
amount  of  rude  force  without  bleeding  ;  nay,  have  to  be  torn 
in  order  to  make  them  bleed.  Then,  too,  the  organs  are 
inclosed  in  a  buccal  pouch,  which  is  filled  through  the  oral 
orifice,  and  emptied  through  a  small  external  opening,  con- 
trolled by  muscles  in  front  of  the  head  fin  ;  so  that  when  the 
animal  respires,  he  can  retain  the  fluid  (air  or  water,  as  the 
case  may  be)  till  completely  exhausted  of  the  oxygen,  then 
discharging  it  through  the  sinuous  gill-opening.  In  fishes, 
however,  the  matter  is  different,  the  gills  being  freely  ex- 
posed to  the  outside  water  by  means  of  the  wide  opercula 
which  simply  cover  them,  and  when  expanded  the  water 
rushes  through  the  organs  in  two  directions,  or  through  the 
gill-opening  as  well  as  the  mouth,  but  more  by  way  of  the 
gill-opening,  all  the  water  escaping,  however,  in  this  direction 
when  the  opercula  are  contracted,  while  the  action  is  much 
more  rapid  than  in  the  eel.  Now,  then,  this  respiratory 
arrangement  in  the  eel  enables  the  animal  to  respire  air  as 
well  as  water,  each  inspiratory  effort  filling  and  distending  the 
buccal  chamber,  causing  it  to  form  a  large  protuberance  upon 
either  side  of  the  head,  which  gradually  collapse  again  as  the 
air  is  slowly  discharged  through  the  small  gill-openings,  when 
they  are  again  inflated  by  inspiratory  effort,  doing  this  several 
times  in  a  minute.     And  by  thus  respiring  air,  the  animal  is 


408  THE   EEL   AMPHIBIOTJ8. 

enabled  to  sustain  its  life  from  six  to  eight  days  out  of 
water,*  so  that  the  animal  can  go  to  distant  waters  over- 
land in  the  night-time,  perhaps,  for  I  am  not  aware  that 
he  has  ever  been  caught  at  it.  Still,  there  can  be  but 
little  doubt  of  the  fact,  as  eels  make  their  appearance  in 
isolated  waters.  At  any  rate,  he  is  certainly  amphibious. 
But  this  is  not  the  route  metamorphosis  takes  in  lung  devel- 
opment, for  the  lungs  are  not  at  the  base  of  the  skull,  but  in 
the  common  visceral  cavity  in  amphibia  and  all  air-breathers,  f 
and  which  is  necessary  for  affording  protection  to  the  delicate 
capillary  network,  at  the  same  time  compelling  absorption 
and  excretion  of  the  gases,  the  lighter  media  of  the  atmo- 
sphere calling  for  this,  that  force  may  be  invoked  for  com- 
pelling the  gases  through  the  membranes  commensurate 
with  the  physiological  requirements.  The  organ  which  is 
the  seat  of  the  changes  for  effecting  these  objects  is  the  air- 
bladder  in  the  fishes,  which  undergoes  progressive  metamor- 
phosis, becoming  more  and  more  vascular,  dividing  and  sub- 
dividing for  increasing  surface  till  all  is  complete  in  the  mam- 
malia. 

In  the  transformations  in  Polypterus,  as  in  Lepidosiren,  the 
ductus  pneumaticus,  which  is  continued  from  the  fore  part  of 
the  posterior  division  of  the  air-bladder  and  opens  into  the 
ventral  part  of  the  beginning  of  the  oesophagus,  is  converted 
into  the  trachea,  with  which  it  is  homologous,  while  the  blad- 
ders undergo  division  and  sub-division  by  longitudinal  and 
transverse  septa  into  cells,  the  work  of  division  for  increasing 
the  vascular  surface  being  carried  through  the  higher  fishes  to 
lepidosiren,  thence  by  amphibia,  reptilia  and  birds  to  the  mam- 
malia, in  which  lung-development  is  complete. 

The  duct  presents  great  variety  in  length,  diameter  and 
place  of  communication  with  the  alimentary  canal.     In  the 

*  I  kept  a  large  fresh-water  eel  in  a  tub  containing  a  thin  stratum  of  water, 
or  sufficient  only  to  cover  the  bottom,  to  enable  the  animal  to  keep  himself 
moist,  but  not  to  respire,  for  seven  days  ;  and  while  he  ceased  to  move  about, 
life  was  not  extinct ;  but  I  finished  it,  deeming  the  experiment  sufficient  for 
the  purpose. 

f  The  intervention  of  a  diaphragm  for  separating  the  viscera  in  the  abdo- 
men from  the  viscera  in  the  chest,  as  occurs  in  mammalia,  has  already  been 
explained  (pp.  84-92,  185-196). 


AIR- BLADDER  THE   NASCENT  LUNGS.  409 

herring,  for  example,  the  posterior  third  of  the  long,  fusiform 
air-bladder  (Fig.  148,  &,  T)  is  connected  with  the' attenuated 
end  of  the  cardiac  division  of  the  stomach.  The  long,  nar- 
row and  flexuous  ductus  pneumaticus  is  continued  from  the 
fore  part  of  the  posterior  division  of  the  air-bladder  in  the 
Cyprinoids,  and  opens  into  the  dorsal  part  of  the  oesopha- 
gus (Fig.  147,  su) ;  the  short,  straight  and  wide  ductus 
pneumaticus  in  the  Lepidosteus  opens  also  into  the  dorsal  part 
of  the  oesophagus,  the  orifice  being  served  by  a  sphincter ;  in 
the  Erythrinus,  the  air-duct  communicates  with  the  side  of 
the  oesophagus  ;  in  Polypterus,  as  in  Lepidosiren,  with  the 
under  or  ventral  part  of  the  beginning  of  the  oesophagus. 

' :  Under  all  its  diversities  of  structure  and  function,  the  homol- 
ogy of  the  swim-bladder  with  the  lungs  is  clearly  traceable ; 
and  finally,  in  those  orders  of  fishes  which  lead  more  directly 
to  the  reptilia,  as,  for  example,  the  salamandroid  Ganoidei  and 
Protopteri,  those  further  modifications  are  superinduced  upon 
the  air-bladder,  by  which  it  becomes  also  analogous  in  function 
to  the  lungs  of  the  air-breathing  Amphibia. 

"The  Lepidosiren  annectens  inhabits  a  part  of  the  river  Gam- 
bia, which  in  the  rainy  season  overflows  extensive  tracts,  that 
are  again  left  dry  in  the  dry  season.  Those  which  do  not  fol- 
low the  retreating  waters  escape  from  the  scorching  rays  of  the 
African  sun  by  burrowing  in  the  mud,  which  is  soon  baked 
hard  above  them ;  but  they  maintain  a  communication  with 
the  air  by  a  small  aperture,  and,  coiling  themselves  up  in  their 
cool  chamber,  clothe  themselves  by  a  layer  of  thick  mucous 
secretion,  and  await,  in  a  torpid  state,  the  return  of  the  rains 
and  the  overflowing  of  the  mud-banks.  The  advent  of  their 
proper  element  wakes  them  into  activity :  they  then  emerge 
from  the  softened  mud,  swim  briskly  about,  feed  voraciously, 
and  propagate. 

' '  The  peculiar  modifications  of  the  gills  and  air-bladder  of  the 
Lepidosiren  are  precisely  those  which  adapt  them  to  the  pecu- 
liar conditions  of  their  existence.  In  the  inactive  state  into 
which  they  are  thrown  by  their  false  position  as  terrestrial 
animals,  the  circulation,  which  would  have  been  liable  to  be 
stopped  had  all  the  branchial  arteries  developed  gills,  as  in 
normal  fishes,  is  carried  on  through  the  two  persistent  primi- 


410  AIR-BLADDER   THE   NASCENT   LUNG?. 

tive  vascula^*  channels.  Whatever  amount  of  respiration  was 
requisite  to  maintain  life  during  the  dry  months  is  effected  in 
the  pulmonary  air-bladders  ;  its  short  and  wide  duct  or  trachea, 
the  oesophageal  orifice  of  which  is  kept  open  by  a  laryngeal 
cartilage,  introduces  the  air  directly  into  the  bladders  ;  the 
blood  transmitted  through  the  branchial  arches  to  the  pul- 
monary arteries  is  distributed  by  their  ramifications  over  the 
cellular  surface  of  the  air-bladders,  and  is  returned  arterialized 
by  the  pdmonary  veins.  A  mixed  venous  and  arterial  blood 
is  thence  distributed  to  the  system,  and  again  to  the  air- 
bladders.  True  arterial  blood  exists  only  in  the  pulmonary 
veins,  and  unmixed  venous  blood  only  in  the  system  of  the 
vena3  cavse  ;  whence  the  necessity,  apparently,  for  that  peculiar 
arrangement  by  which  the  arterial  blood  is  conveyed  directly 
to  the  ventricle  by  the  pulmonary  vein.  When  the  Lepidosiren 
resumes  its  true  position  as  a  fish,  the  branchial  circulation  is 
vigorously  resumed,  a  larger  proportion  of  arterialized  blood 
enters  the  aorta,  and  both  the  nervous  and  muscular  systems 
receive  the  additional  stimulus  and  support  requisite  for  the 
maintenance  of  their  energetic  actions  "  (Owen). 

And  that  this  is  the  true  genesis  of  the  lungs  is  fully  proven 
by  the  following  incontrovertible  facts,  namely  : 

1.  In  Batrachia  this  condition  of  double -breathing  is  seen 
only  in  the  immature  animal,  or  tadpole,  which  in  passing 
through  the  consecutive  changes  rids  itself  of  the  branchiae 
and  tail ;  also  involving  important  correlated  changes  in  other 
organs,  notably,  heart,  lungs,  vascular  system,  etc. 

2.  In  embryogenesis  of  warm-blooded  animals,  in  which  the 
whole  thing  repeats  itself,  the  lungs  is  but  a  differentiated 
diverticulum  of  the  intestinal  canal,  the  same  as  the  air-bladder, 
the  point  of  in-folding  of  the  membranous  layers  so  as  to 
form  the  trachea  and  lungs  being  at  the  anterior  and  upper 
end  of  the  gullet,  corresponding  with  the  pneumatic  duct 
and  air-bladder  in  the  higher  fishes,  Lepidosiren,  Axolotl, 
Batrachia  and  all  reptilia.  Furthermore,  the  pulmonic,  vas- 
cular and  cardiac  changes  are  easily  traceable  through  all  the 
subsequent  changes  which  occur  In  the  Axolotl  (Fig.  169), 
in  which  the  respiratory  and  circulatory  organs  are  substan- 
tially the  same  as  in  the  tadpole,  we  have,  for  example,  the 


DIFFERENTIATION"   OF   LEFT  AURICLE.  411 

fourth  pair  of  branchial  arches  (5)  continued  on  into  the  air- 
Madders  or  nascent  lungs,  the  upper  three  passing,  as  usual, 
into  the  long  fibriated  branchiae  projecting  from  the  sides  of 
the  head  ;  the  pulmonary  veins,  one  for  either  side,  returning 
the  blood  to  the  small  left  auricle  inclosed  in  the  walls  of  the 
large  right  auricle,  as  there  is  no  projection  externally  in- 
dicating its  existence  (which  at  first  led  to  the  impression  that 
but  one  auricle  existed  in  the  double-breathers  or  perenni- 
branchiata    amphibious) ;    the    branchiae    gradually  become 
dim  inished  in  size,  the  vessels,   of  course,  shrinking  in  pro- 
portion,  while  the  lungs  are  progressively  more  and  more 
developed,  the  pulmonary  artery  and  vein  expanding  in  pro- 
portion, till  at  last  we  have  all  the  vascular,  pulmonic  and 
cardiac  changes  that    obtain  in   the  frog  (Fig.    170).      The 
branchial  arches   are   absorbed   in  proportion  as  the  circu- 
lation becomes  modified,  their  atrophy  depending  upon  the 
changes  which  take  place  in  the  course  of  the  blood,  owing  to 
the  dilatation  of  the  anastomotic  vessels  and  enlargement 
of  the  pulmonary  artery.     In  Lepidosiren,  which  is  the  lowest 
of  the  perennibranchiates,  the  heart  resembles  that  of  a  fish, 
consisting  of  a  single  auricle,  ventricle  and  bulbus  arteriosus. 
The  vena  cava,  bringing  the  vitiated  blood  from  the  system, 
terminates  at  once  in  the  auricle,  while  the  pulmonary  vein 
passes  along  as  far  as  the  auriculo-ventricular  opening,  where 
it  empties  its  contents  into  the  ventricle  by  a  distinct  orifice, 
guarded  by  a  cartilaginous  valvular  tubercle  (Owen). 

The  following  brief  summary  by  Owen*  indicates  the  cardiac 
changes  in  the  amphibia : 

"The  distinction  of  the  pulmonary  from  the  systemic  auricle, 
first  observed  in  Siren,  has  been  since  determined  in  Meno- 
branchus,  f  Axolotes,  J  AmpMuma,  §  and  Menopoma.  In  Pro- 
teus, in  which  some  of  the  blood  of  the  puny  lungs  is  conveyed 
to  systemic  veins,  the  auricular  septum  is  not  complete,  accord- 
ing to  Hurd.  1    In  Ampliiuma,  the  auricle  is  smaller  and  less 

*Ibid.,  pp.  506-507. 

f  Meyer.     Analekten  fur  Vergleichende  Anatomie,  4to,  p.  73.     1835. 

%  Calori.     Sull'  Anatomia  dell'  Axolotis  Commentario,  4to,  p.  45. 

§Ibid.,  p.  215. 

J]  John  Hunter.     On  the  Blood  and  Inflammation.     4to,  p.  258.     1794. 


412 


DIFFKRENIIATION   OF   LEFT  AURICLE. 


fimbriated  than  in  Siren.  The  ventricle  is  similarly  connected 
to  the  pericardium  by  the  apex,  as  well  as  by  the  artery.  This 
formsahalf-spiial  turn  at  its  origin,  and  dilates  into  a  broader 
and  shorter  bulb  than  in  siren.  In  Menopoma,  they  are  still 
more  reduced  in  size,  and  lie.  as  in  Salamandra*  (Pig-  171,  a), 
when  nndistended,  to  the  lef  r  of  the  ventricle ;  their  outer  surface, 


Fig.  169. — Respiratory  and  Circulatory  Organs  in  Prbteiis  Mexicanus  (Axolotl). — Cyclo- 
paedia of  Anatomy,  Art.  Respiration  (Supplement).  Allen  Thomson.  A,  truncus 
arteriosus ;  B,  the  three  upper  pairs  of  branchial  arteries ;  P,  P,  the  lower  pair 
proceeding  to  the  lungs  or  pulmonic  arteries  ;  b,  branchial  veins  ;  v,  pulmonary  veins  ; 
V,  vena  cava  ;  I,  hepatic  vein  ;  H,  ventricle  of  the  heart,  surmounted  by  the  large 
right  auricle. 

as  in  Menobranchus,  is  entire.  The  ventricle  is  of  a  flattened, 
triangular  form  ;  its  cavity  is  occupied  by  the  loose,  fasciculate 
muscular  structure,  through  which  the  blood  filters,  as  through 
a  sponge,  from  the  small  contiguous  auricular  apertures,  each 
of  which  has  a  simple  valve,  to   the    'ostium    arteriosum.' 

*  Owen,  Fig.  333. 


DIFFERENTIATION-    OF   LEFT   AURICLE. 


413 


The  artery  inclines,  with  a  slight  twist,  to  the  left,  and  swells 
into  a  spherical  bulb.  The  valves  are  confined  to  the  narrower 
part,  and  are  in  two  transverse  rows,  four  in  each  row,  each 
valve  of  a  conical  shape,  pointing  forward.*  The  first  row  is 
just  above  the  ostium  ;  the  second  is  half-way  between  this 
and  the  bulb. 

' '  The  pulmonic  auricle  augments  in  size  with  the  more  ex- 


Fig.  170. — Respiratory  and  Circulatory  Organs  in  the  Frog,  showing  increase  in 
pulmonic  and  cardiac  development. — Williams,  a,  Vena  cava  ;  6,  right  auricle  ; 
c,  pulmonary  veins  ;  d,  sinus  ;  i,  left  auricle  ;  e,  ventricle  ;  /,  Truncus  arteriosus  ; 
g,  right  carotid  artery  ;  k,  left  carotid  ;  h,  vessel  to  anterior  extremity  I,  vessels  to 
mandibular  region  and  face  ;  m,  abdominal  aorta,  formed  by  the  junction  of  the 
two  aortse. 

elusive  share  taken  by  the  lungs  in  respiration ;  but  the 
auricular  part  of  the  heart  shows  hardly  any  outward  sign  of 
its  division  in  batrachians.  It  is  small,  smooth,  and  situ- 
ated to  the  left,  and  in  advance  of  the  ventricle,  in  newts  and 
salamanders  (Fig.  17 1). 

"  In  frogs  and  toads,  the  auricle  is  applied  to  the  base  of  the 
ventricle,  and  to  the  back  and  side  of  the  aorta  and  its  bulb. 
The  ventricle,  usually  of  a  more  rounded  form,  is  occupied  by 

*  Catalogue  of  the  Physiological  Series  in  the  Museum  of  the  Royal  College 
of  Surgeons.  4to,  5  vols.  1833-1840.  Second  ed.,  Vol.  I.,  1852.  Richard 
Owen. 


414  INCREASING  DIFFERENTIATION 

the  muscular  fasciculi,  except  at  a  small  part  between  the 
auriculo-ventricular  and  aortal  orifices.  The  bulbus  arteriosus 
is  incompletely  divided  by  opposite  longitudinal  folds,  the 
margins  of  which  meet,  but  remain  free."     Italics  are  added. 

It  will  thus  be  seen  that  there  is  increasing  differentiation  in 
the  cardiac  chambers  coincident  with  lung  development,  in  the 
development  of  a  left  auricle  especially,  which  comes  into 
existence  only  in  connection  with  lung  elaboration,  and  as 
necessary  to  it  for  receiving  and  transmitting  the  aerated  blood 
to  the  systemic  vessels  or  aortse  for  distributing  it  through  the 
body  ;  while  with  increasing  differentiation  in  the  lungs  with 
progress  in  development  a  left  ventricle  also,  by  segmentation 
or  division  of  the  common  ventricle  by  means  of  a  membranous 
partition,  which  is  gradually  differentiated  from  the  walls  so 
as  to  form  the  left  ventricle,  into  which  the  left  auricle  dis- 
charges its  blood,  thereby  completely  isolating  the  blood  com- 
ing from  the  pulmonic  capillaries  from  the  venous  blood  com- 
ing into  the  right  side  of  the  heart,  sending  it  out  through  the 
aortal  system,  which  now  communicates  with  the  left  ven- 
tricular chamber  only  ;  while  the  differentiation  of  this 
partition- wall  is  also  easily  traceable,  being  perfected  in  croco- 
dilia,  only  that  there  is  left  a  communication  between  the 
aortse  above  the  heart,  which  subserves  useful  purpose  during 
periods  of  long  submergence,  allowing  the  venous  blood  to 
pass  from  the  left  to  the  right  aorta.     Of  this  more,  further  on. 

The  following  from  the  greatest  of  living  naturalists,  per- 
haps, completes  the  description  of  the  cardiac  and  vascular 
changes  in  perennibranchiates  and  batrachians  :* 

"  The  heart  presents  two  auricles,  a  single  ventricle  and  a 
buibus  arteriosus.  A  venous  sinus,  the  walls  of  which  are 
rhythmically  contractile,  receives  the  venous  blood  from  the 
body,  and  opens  into  the  right  auricle.  In  Proteus,  Meno- 
branchus  and  Siren,  the  septum  of  the  auricles  is  less  com- 
plete than  in  the  other  Amphibia.  The  left  auricle  is  much 
smaller  than  the  right,  and  a  single  pulmonary  vein  opens  into 
it.     The  interior  of  the  ventricle  is  more  like  a  sponge  than  a 

*  Anatomy  of  Vertebrate  Animals.  By  Thomas  H.  Huxley,  LL.D.,  F.  R.  S., 
author  of  "Lay  Sermons,"  "Man's  Place  in  Nature,"  "Origin  of  Species," 
etc.,  etc. 


IN   PERENKTliRANi  HIATES. 


415 


chamber  with  well-defined  parietes.  The  walls  of  the  long 
bulbus  arteriosus  contain  striated  muscular  fibres,  and  are 
rhythmically  contractile.  Yalves  are  sometimes  placed  at 
each  end  of  it,  and  it  may  be  imperfectly  divided  into  two 
cavities  by  an  incomplete  longitudinal  partition.  1 1  terminates, 
upon  each  side,  in  either  three  or  four  trunks,  which  ascend 
upon  the  branchial  arches.  The  most  anterior  of  these  trunks 
give  off  the  carotid  arteries,  the  most  posterior  the  pulmonary 
arteries,  and  arteries  to  the  integument ;  the  middle  trunks 
form  the  principal  roots  of  the  dorsal  aorta. 


Pig.  171. — Vascular  System  and  Hyo-Branchial  Apparatus,  Salamander. 

"  In  Proteus,  where  there  are  three  branchial  arches,  the  buib 
of  the  aorta  splits  into  two  trunks ;  each  of  these  divides  at 
first  into  two  branches,  and  then  the  posterior  branch,  on  each 
side,  again  subdivides  into  two  others.  Thus,  three  pairs  of 
aortic  trunks  are  formed,  which  ascend  upon  the  branchial 
arches.  The  two  anterior  pairs  of  aortic  trunks  pass  directly 
into  the  roots  of  the  dorsal  aorta,  but  each  gives  off  a  vessel 
which  enters  one  of  the  external  gills,  the  blood  from  which  is 
brought  by  an  efferent  canal  into  a  higher  part  of  the  same 
aortic  arch.  The  third  aortic  trunk,  on  each  side,  is  inter- 
rupted, its  lower  part  becoming  the  branchial  artery  of  a  gill- 
tuft.  The  blood  is  carried  out  of  this  branchia  by  a  venous 
trunk,  which  opens  into  the  root  of  the  dorsal  aorta,  and  is,  in 
Teality,  merely  the  upper  part  of  the  third  aortic  trunk.     The 


416        CARDIAC   DEVELOPMENT   ADVANCED   IN   THE   FROCK 

facts  may  be  expressed  in  another  way,  by  saying  that  the 
bases  of  the  branchial  artery  and  vein  anastomose  in  the  iirst 
two  gills,  but  not  in  the  third. 

' l  The  adult  Axolotl  (Siredon)  has  four  pairs  of  aortic  trunks  ; 
the  hindermost  pair  gives  off  the  pulmonary  arteries,  the  three 
next  supply  the  external  branchiae,  and  the  anterior  trunk 
passes  above  into  an  artery  which  divides  into  hyoidean  and 
carotid  branches. 

'  In  Salamnndra  there  are  four  pairs  of  aortic  trunks  in  the 
adult,  but  the  upper  moiety  of  the  first,  on  each  side,  is  oblit- 
erated, and  remains  as  a  mere  ductus  Botdlli.  The  fourth 
trunk  gives  off  the  pulmonary  artery ;  some  twigs  for  the 
oesophagus  and  a  few  cardiac  branches  next  arise  from  it ;  and 
it  then  unites  with  the  second  and  third  to  form  the  root  of  the 
dorsal  aorta.  The  basal  moiety  of  the  first  trunk  enlarges  at 
its  extremity,  close  to  the  angle  of  the  mandible,  into  a  spongy 
organ,  the  carotid  gland,  from  which  the  carotid  artery,  and 
that  for  the  supply  of  the  hyoidean  and  oral  regions,  are  given 
off. 

"  In  the  adult  frog,  the  aortic  bulb  is  separated  by  an  incom- 
plete longitudinal  septum  into  two  passages,  and,  at  its  extrem- 
ity, divides  into  two  trunks,  each  of  which  is  partitioned 
internally  into  three  passages.  The  middle,  or  systemic,  pas- 
sage passes  directly  into  a  trunk,  which  unites  with  its  fellow 
beneath  the  spinal  column  into  the  dorsal  aorta.  The  anterior, 
or  carotid,  passage  ends,  as  in  Salamandra,  in  a  carotid  gland 
and  ductus  Botdlli ;  carotid,  hyoidean  and  oral  branches 
being  given  off  from  the  former.  The  hindermost,  or  pulmo- 
cutaneous,  passage  ends  in  the  pulmonary  and  the  cutaneous 
arteries,  the  anastomoses  of  these  with  the  roots  of  the 
dorsal  aorta  being  obliterated.  The  middle  pair  of  aortic 
trunks  thus  exclusively  constitute  the  origins  of  the  dorsal 
aorta,  and  are  the  permanent  aortic  arches.  The  right  aortic 
arch  is  wider  than  the  left,  especially  toward  their  junction  ; 
as  the  left  gives  off,  just  before  this  point,  a  large  coeliaco- 
mesenteric  artery  to  the  abdominal  viscera.  Each  aortic  arch 
gives  off  the  subclavian  and  vertebral  arteries  of  its  side.  Only 
venous  blood  passes  into  the  pulmonary  arteries  of  a  frog; 
while  mixed  blood  enters  the  aortic  arches,  and  is  of  a  brighter 


CAEDIAC   DEVELOPMENT   ADVANCED   IN   THE  FROG.        417 

arterial  hue  at  the  end  than  at  the  beginning  of  the  systole 
The  blood  in  the  carotid  passages  is  always  bright.  The 
mechanical  arrangements  by  which  this  is  brought  about  have 
been  beautifully  analyzed  by  Briicke,  who  shows,  first,  that 
the  spongy  interior  of  the  ventricle  contains,  in  its  base,  a 
transversely-elongated  cavity,  into  which  the  auricles  open, 
and  which,  by  its  right  extremity,  communicates  with  the  ven- 
tricular opening  of  the  aortic  bulb  ;  secondly,  that  the  aortic 
bulb  is  imperfectly  divided  by  a  longitudinal  septum,  the 
upper  left  edge  of  which  is  attached,  while  its  lower  right  edge 
is  free  ;  thirdly,  that  of  the  two  passages  into  which  the  aortic 
bulb  is  thus  divided,  the  one  on  the  right  side  of  the  septum 
ends  in  a  chamber,  in  which  the  carotid  and  systemic  passages 
commence,  while  that  on  the  left  side  similarly  leads  to  the 
entrance  to  the  pulmo-cutaneous  passages  ;  fourthly,  that  the 
carotid  gland,  in  which  the  carotid  passage  ends,  presents  a 
mechanical  obstacle  to  the  flow  of  the  blood  through  it ;  fifthly, 
that  there  is  a  valvular  fold  open  toward  the  heart,  in  each 
systemic  passage,  which  also  offers  a  certain  amount  of  me- 
chanical resistance  to  the  blood ;  and,  sixthly,  that  after  the 
blood  has  begun  to  flow  through  the  bulb,  it  will  gradually 
force  the  septum  over  to  the  left  side,  and  so  impede  the  flow 
into  the  pulmo-cutaneous  passage. 

"Thus,  when  the  auricular  systole  takes  place,  the  right 
auricle  sends  its  venous  blood  into  that  division  of  the  ven- 
tricular cavity  which  lies  nearest  the  opening  of  the  bulb  ;  and 
when  the  ventricle  contracts,  the  blood  first  driven  into  the 
bulb  is  wholly  venous.  This  blood  fills  the  passages  on  both 
sides  of  the  septum,  but  finds  a  very  much  greater  resistance  to 
its  exit  on  the  right  than  on  the  left  side.  It  therefore  flows, 
at  first,  exclusively  into  the  left  division,  and  makes  its  way 
through  the  short  pulmonary  arteries  into  the  lungs.  But,  as 
the  pulmonary  vessels  fill,  the  pressure  on  the  two  sides  of  the 
septum  becomes  equalized,  and  the  systemic  passages,  which 
offer  the  next  least  resistance,  fill  with  blood,  which  is  now 
mixed,  as  it  comes  from  the  middle  of  the  ventricle  Next,  the 
septum,  being  driven  over  to  the  left  side,  prevents  any  more 
blood  from  going  into  the  pulmo-cutaneous  passage.  At  the 
end  of  the  systole,  the  blood  driven  out  by  the  ventricle  is 


416     PECl'I.IARITIES   IN   PERENNIBRANCH'ATE   RESPIRATION. 

almost  wholly  that  of  the  left  auricle  ;  and,  by  this  time,  the 
resistance  in  the  systemic  is  as  great  as  that  in  the  carotid  pas- 
sages. Hence,  the  latter  fill  and  send  arterialized  blood  to  the 
head. 

"The  organs  of  respiration  of  the  Amphibia,  in  the  adult 
state,  are  either  external  branchiae,  combined  with  lungs,  as  in 
the  perennibranchiate  Urodela;  or  lungs  only,  as  in  the  other 
Urodela,  the  Batrachia,  the  Gymnophiona,  and,  probably,  the 
majority  of  the  Labyrinthodonta. 

"In  the  perennibranchiate  Urodela,  the  branchial  arches  (or 
some  of  them)  are  separated  by  open  clefts  (the  number  of 
which  varies  from  four  to  two),  throughout  life,  and  three 
branched  gills  are  continued  by  single  stems  into  the  integu- 
ment, at  the  dorsal  ends  of  the  branchial  arches.  An  opercular 
fold  of  the  integument,  in  front  of  the  gill-clefts,  attains  a  con- 
siderable size  in  Siredon,  but  does  not  cover  the  gills.  The 
branchial  arches  themselves  bearno  branchial  filaments.  Other 
Urodela  are  devoid  of  external  gills,  but  (as  is  the  case  in 
Menopoma  and  Amphiuma)  present  one  or  two  small  gill- 
clefts  on  each  side  of  the  neck,  and  are  thence  called  Derotre- 
mata.  The  rest  of  the  Urodela,  and  all  the  Batrachia  and 
Gymnophiona,  are  devoid  of  both  external  gills  and  gill-clefts, 
in  the  adult  state. 

"  In  all  the  Amphibia,  a  glottis,  placed  on  the  ventral  wall  of 
the  oesophagus,  opens  into  a  short  laryngo- tracheal  chamber 
with  which  two  pulmonary  sacs  are  connected,  either  directly 
or  by  the  intermediation  of  bronchi  (as  in  the  Aglossa),  or  by 
a  trachea  (as  in  the  Gymnophiona).  The  walls  of  the  pulmo- 
nary sacs  are  more  or  less  sacculated.  In  most  Amphibia  the 
lungs  are  equal  in  size  ;  but  in  the  snake-like  Gymnophiona 
the  right  is  much  smaller  than  the  left.  In  Proteus,  the  pul- 
monary blood  is  not  all  returned  to  the  heart,  some  of  it  enter- 
ing the  veins  of  the  trunk.  Aerial  respiration  is  effected,  in  the 
Amphibia,  by  pumping  the  air  from  the  oral  cavity  into  the 
lungs.  To  this  end  the  mouth  is  kept  shut,  and  ingress  and 
egress  to  the  air  is  given  by  the  nasal  passages,  which  always 
open  immediately  behind  the  vomers,  at  the  anterior  part  of 
the  roof  of  the  mouth.  These  passages  being  open,  and  the 
hyoidean  apparatus  depressed,  the  air  fills  the  cavity  of  the 


EESPI RATION   IN   CHELONTA.  419 

mouth.  The  external  nostrils  are  then  shut,  and,  the  hyoidean 
apparatus  being  raised,  the  air  is  forced,  through  the  open 
glottis,  into  the  lungs." 

Professor  Huxley,  in  common  with  all  naturalists,  conceives 
that  the  frog  respires  only  by  pumping  the  air  into  the  lungs 
by  means  of  the  throat-apparatus,  that  action  of  which  is  very 
conspicuous  ;  and  while  it  undoubtedly  is  of  great  assistance, 
that  it  is  not  fundamental  in  respiration  I  have  fully  demon- 
strated by  physiological  experiment  (pp.  61-68).  In  c/ielonia, 
however,  for  special  reasons  that  obtain  in  them,  it  is  sub- 
stantially but  not  entirely  correct,  and  since  it  has  direct 
bearing  upon  cardiac  development,  we  shall  proceed  to  con- 
sider it  somewhat  at  length,  bringing  out  the  special  circum- 
stances in  the  anatomy  of  the  animal  for  throwing  light  upon 
its  mode  of  respiration.  We  have  seen  that  the  lungs  and 
body-envelope  expand  and  contract  together  or  simultaneously, 
regularly  and  rhythmically,  in  order  to  produce  afflux  and 
efflux  of  the  fluids  in  the  alveolar  chambers  for  respiratory 
purposes,  and  must  do  so  in  the  very  nature  of  things,  and  it 
remains  to  explain  the  exceptional  conditions  which  obtain  in 
chelonia,  and  how  respiration  and  circulation  are  compelled 
to  be  in  correspondence  with  the  activities,  notwithstanding 
the  seeming  drawbacks  and  apparent  obstacles.  In  the  first 
place,  we  have  to  note  the  house-like  body  formed  by  the 
carapace  (above)  and  plastron  (below)  (Fig.  172),  containing  the 
viscera,  but  with  openings  anteriorly  and  posteriorly  for  the 
head  and  extremities,  which  work  out-and-in  the  excavation 
during  extension  and  retraction,  thereby  producing  corre- 
sponding changes  of  pressure  in  the  cavity  and  lungs  during 
locomotion  ;  and  the  more  rapid  this  is  made,  the  more  effective 
it  should  be.  By  reason  of  the  unyielding  floor  of  support  to 
the  viscera  in  the  plastron,  expansion  is  impossible  in  this 
direction,  so  that  the  animal  gains  nothing  from  the  action  of 
gravitation,  as  in  the  other  animals,  but  ample  arrangements 
obtain  nevertheless  for  compelling  respiration  to  be  in  corre- 
spondence with  the  activities  ;  otherwise  these  could  not  for 
obvious  reasons  be  produced.  And  by  first  extending,  then 
retracting,  the  extremities  by  means  of  the  special  muscles 
which  have  their  points  of  origin  toward  the  central  portions 


420  RESPIRATION   II?    <  IIllLoNIA. 

of  the  floor  or  plastron,  scapular  and  pelvic  bones  (FigB. 
172,  192),  corresponding  increase  and  diminution  of  the  vis- 
ceral area  are  made  inevitable.  But  to  this  again  must  be 
added  the  action  in  the  special  muscles  which  apply  Tor  effect- 
ing changes  of  pressure  in  the  lungs,  notably  :  1.  The  trans- 
versus  abdominis  (Fig.  172,41),  and  obliquus  abdominis  (40) 
(one  for  either  side),  which  are  homologous  with  those  in  the 
higher  animals,  closing  in  the  posterior  end  upon  the  sides, 
the  muscles  of  the  posterior  extremities  with  their  points  of 
origin  toward  the  central  portions  of  the  carapace  (45,  95,  9tf, 
103,  5G,  101,  48,  etc.),  closing  the  intervening  portions;  while 
anteriorly  we  have  the  muscles  of  the  scapulae  and  anterior  ex- 
tremities, together  with  the  diaphragmaticus  (42),  serratus  mag- 
nus  (57),  closing  in  the  antero-lateral  regions,  the  head  and  cer- 
vical muscles  the  intervening  portions.  Now,  then,  in  order  to 
study  the  action  in  these  muscles  for  determining  which  are  the 
more  effective  in  respiration,  the  anterior  or  posterior,  we  must 
place  the  animal  in  a  vessel  of  water  and  note  the  respirations 
and  the  effect  they  produce.  For  example,  when  the  animal  is 
floating  lazily  upon  the  water,  the  limbs  projecting  out  to  the 
full  extent,  over  the  edge  of  the  plastron,  the  head  and  tail  also 
extended,  it  will  be  seen  that  the  top  of  the  carapace  project- 
ing partially  above  the  water,  will  every  now  and  then  leap 
suddenly  upward,  then  settle  down  again  as  suddenly  to  the 
former  level,  and  if  at  the  same  time  the  "flanks  "  be  observed, 
they  will  be  seen  to  project  with  the  former  and  to  retract  with 
the  latter,  showing  conclusively  respiratory  movement ;  the 
former  corresponding  with  inspiration,  the  latter  with  expira- 
tion, of  which  there  can  be  no  doubt,  since  the  effect  of  the  one  is 
to  increase  the  amount  of  air  in  the  lungs,  the  other  to  diminish 
it ;  the  one  causing  the  upward,  the  other  the  downward 
movement,  from  the  diminution  and  augmentation  in  body- 
density  ;  otherwise  is  inexplicable. 

Furthermore,  these  muscles  in  the  flanks  are  incurvated 
inward,  projecting  into  the  excavation  like  the  diaphragm  in 
the  chest-excavation,  the  convexity  toward  the  lungs  ;  hence, 
when  they  contract,  tend  to  pull  away  from  the  lungs,  thereby 
reducing  pressure  in  the  excavation  the  same  as  in  the  other 
case,    the   air  rushing    into    the    locality   simultaneously  to 


KESPIKATIOIST   IIST    CHELOISTIA. 


421 


equalize  pressure,  the  lungs,  of  course,  acting  in  concert 
with  the  respiratory  muscles  as  with  the  diaphragm.  As 
will  be  seen,  the  lungs  (Fig.  173,  z,  z)  are  very  large,  cov- 
ering the  entire  viscera  upon  the  dorsal  surface ;  and  while 
numerous  longitudinal  and  transverse  septa  divide  and  sub- 
divide the  interior  (Fig.  174),  the  chambers  are  comparatively 
large  and  freely  communicate  with  each  other  and  the  primi- 
tive division  of  the  trachea  (a),  the  bristles  indicating  this  cir- 
cumstance ;  they  are  not  bound  down  or  fastened  by  the  lining 
membrane  so  as  to  limit  the  action,  but  are  free  to  move,  glid- 
ing readily  to  and  fro  upon  the  viscera  in  respiration,  expand- 


ing. 172.—  Showing  the  Myology  of  the  Tortoise  (Emys  Europcea),  with  the  special 
muscles  relating  to  respiration. — Bojanus.  Anatome  Testudinis  Europaea.  Fol. 
1819-1821.  41,  Transversus  abdominis ;  40/7 ',  o'bliquus  abdominis ;  42,  diaphrag- 
maticus  ;  57,  serratus  magnus. 

ing  laterally  as  well  as  longitudinally,  the  copious  secretions 
in  the  parts  enabling  this  action,  as  also  the  actions  appertain- 
ing to  the  viscera  themselves  in  connection  with  their  special 
f  unctions,  or  the  same  as  obtains  in  warm-blooded  animals. 
Finally,  for  effecting  coordination  with  respiration,  we  have 
the  nerves  proceeding  to  the  several  parts  from  the  spinal 
medulla  (Fig.  192),  inclusive  of  the  heart  and  vessels,  all  the 
parts  being  fully  coordinated  for  producing  a  balance  in  force, 
the  same  as  in  all  the  animals,  and  must  be  so,  in  the  nature  of 
things  ;  the  thoracic  duct  (c)  coursing  upon  the  aortse  to  reach 
the  chyle  receptacles  (5,  b)  and  veins  at  the  root  of  the  neck  be- 
ing alike  included,  so  that  when  the  animal  locomotes,  ner- 
vous force  throbs  over  all  the  structures  for  increasing  circu- 


422 


KKSPIKATIO.N    IN   CIIELoNlA. 


lation  correspondingly  for  maintaining  a  balance  in  the  organ- 
ism ;  otherwise  impossible. 

This  gives  ns  the  respiratory  action.  Now,  then,  in  regard 
to  the  throat-apparatus  and  the  special  role  it  performs  in 
respiration,  the  importance  of  which  it  would  be  difficult  to 


Fig.  173. — The  Viscera  in  the  Tortoise  (Emys  Europcea),  as  viewed  from  above. — 
Bojanus.  Z,  Z,  lungs  ;  I,  liver  ;  7,  oesophagus  ;  E,  trachea  ;  F,  F,  axillary  arteries; 
6,  chyle  receptacle  at  the  root  of  the  neck,  one  for  either  side  ;  i,  jugular  vein  ;  h, 
brachial  vein  ;  X,  urinary  bladder  ;  o,  kidney  ;  H,  urethra  ;  V,  V,  intestines  ;  U,  U, 
anal  bursa}  ;  C,  thoracic  duct. 

overestimate,  being  essential  to  life.  Briefly,  it  consists  as  fol- 
lows :  The  body  of  the  hyoid  bone  projects  from  its  sides  three 
pairs  of  long,  curved,  rib-like  processes  (Fig.  192,  d),  which  are 
operated  by  muscles  connecting  with  the  mandible  and  base  of 
the  skull,  similar  to  what  obtains  in  the  frog,  but  much  more 
powerful,  the  whole  inclosed  by  the  muscular  sheath  {latissi- 


KESPI..ATION   IJST    CHELONIA-  423 

mus  colli  (Fig.  172,  21,  25)  expanded  over  it,  connecting  with  the 
mandible  and  base  of  the  skull,  imparting  the  rotund  appear- 
ance and  fullness  upon  the  sides  and  inferior  aspects,  the 
underlying  hyoid -framework  pushing  out  the  soft  structures. 
As  in  the  frog,  the  pumping  action  is  very  rapid,  and  goes 


Mg.  174.— Lung  of  a  Tortoise  (Emys  Europcea).  — Bojanus.  The  bristles  indicate  the 
free  passage  of  air  through  the  lung  structure,  the  chambers  all  freely  communicat- 
ing with  each  other  and  the  trachea  (a) ,  or  bronchus. 

on  all  the  while,  in  the  water  as  well  as  out  of  it,  which  led 
John  Hunter  to  infer  that  it  does  not  relate  to  respiration, 
which,  in  view  of  the  extraordinary  acuteness  of  perception 
which  characterized  that  grand  pioneer  in  natural  history,  is  a 
little  remarkable,  and  explicable  upon  the  ground  simply  that 
he  was  not  guided  by  this  fundamental  law  underlying  animal 
structure  and  function,  but  seeking  out  the  history  in  develop- 


424  RESPIRATION   IN    CHELONIA 

ment  itself ;  performing  enormous  labors  with  a  singleness  of 
purpose  and  devotion  to  make  him  memorable  as  long  as  rea- 
son lives  and  noble  deeds  are  exemplars  for  men.  The  tor- 
toise, as  well  as  the  frog,  when  it  goes  beneath  the  water, 
must  bottle  up  air  in  the  lungs  and  air-pas?ages  by  means  of 
the  lingual  valve  applied  to  the  posterior  nares,  pumping  it 
backward  and  forward  in  the  lungs  till  the  oxygen  is  con- 
sumed (p.  66),  when  it  must  return  to  the  surface  for  a  fresh 
supply.  Being  an  air-breather,  it  is  necessarily  compelled  to 
this  course,  taking  a  fresh  supply  down  with  it  every  time  it 
goes.     All  of  which  is  easily  understood. 

Now,  then,  in  order  to  ascertain  the  relative  amount  of 
work  the  apparatus  performs  in  respiration,  1  had  recourse 
to  the  same  experiment  performed  in  the  case  of  the  frog, 
namely,  excision  of  a»  portion  of  the  floor  of  the  mouth. 
Using  a  tenaculum  hooked  under  the  mandible,  I  forcibly 
drew  out  the  head  and  neck,  and  with  a  pair  of  sharp-pointed 
scissors,  one  blade  of  which  was  pushed  through  the  structures 
to  the  interior  of  the  mouth,  I  succeed  in  excising  a  portion 
of  the  floor  sufficient  for  the  purpose,  cutting  through  two  of 
the  ribs  before  accomplishing  it,  the  animal  making  a  noise 
as  if  in  pain.  I  was  sorry  it  had  not  been  chloroformed  ;  but 
being  deeply  absorbed,  it  escaped  me  at  the  time  The  effect 
of  the  operation  was  very  decided,  the  animal  rapidly  becom- 
ing limp  and  motionless,  but  every  now  and  then  making  a 
peculiar,  distressful  noise,  as  if  in  pain — sharp,  quick,  vibra- 
tory sounds,  very  shrill,  plaintive,  and  full  of  pathos  to  me.  It 
touched  me  to  the  quick,  but  too  late  for  repentance.  At  12 
p.  m.,  ten  hours  after  the  operation,  when  I  retired  to  rest,  life 
appeared  extinct ;  yes,  to  all  appearance  it  was  dead.  But 
I  was  suddenly  startled  out  of  sleep  at  1  a.  m.  by  a 
shrill  sound,  which  I  at  once  recognized  as  the  peculiar  cry 
of  distress  coming  from  the  tortoise  ;  followed  by  the  sound  of 
something  falling  down,  which  I  could  not  make  out,  as  I  had 
left  it  upon  the  floor  in  the  room  adjacent.  Springing  out  of 
bed,  for  investigation,  I  found  the  animal  upon  its  back  in  a 
corner  of  the  room,  where  it  had  crawled,  and,  I  suppose, 
struggling  upon  its  hind  limbs  against  the  wall  for  air,  fell 
over  dead,  giving  vent  to  the  expiring  cry  as  it  did  so.     Tak- 


RESPIRATION   IN   CHELOMA.  425 

ing  it  up  in  my  hands,  the  head  and  limbs  swung  loosely 
by  their  attachments  :  it  was  dead.  Ah  me  !  the  sorrow.  I 
was  overworking  at  the  time  (I  fear  me,  I  have  done  that  too 
long),  and  perhaps  it  had  something  to  do  with  the  acuteness 
of  my  senses  and  feelings,  which  was  very  likely  ;  but  have 
never  felt  like  repeating  the  experiment.  JSTein,  das  ist 
genug  /*  Upon  the  following  day,  I  made  a  careful  dissec- 
tion to  ascertain  how  the  loud  vocal  resonance  had  been  pro- 
duced, since  it  involved  forcible  com|3ression  of  the  lungs 
for  rushing  the  air  through  the  vocal'  cords,  which  made  it 
manifest  that  expiration  is  much  more  energetic  in  the  tor- 
toise than  inspiration.  The  explanation  is  easy.  Thus,  the 
diaphragmatici  (one  upon  either  side)  (Fig.  172,  4:)  closely 
embrace  the  lungs,  anteriorly  and  laterally,  and,  by  contract- 
ing, must  inevitably  compress  the  organs.  There  is  no  doubt, 
then,  this  was  the  explanation  of  the  phenomenon.  It  was 
an  error  in  Bojanus  to  have  named  this  muscle  the  diaphragm- 
aticus,  tending  to  misdirection ;  for  there  is  nothing  in  the 
physiology  of  the  tortoise,  nor  in  any  of  the  animals  antece- 
dent to  the  mammalia,  calling  for  it,  the  differentiation  of  a 
diaphragm  being  correlated  with  the  changes  in  the  intes- 
tines, converting  them  into  large  air-chambers  for  increasing 
the  digestive  and  absorptive  processes,  etc.,  for  restraining 
them  so  as  not  to  interfere  with  respiration,  but  rather  ener- 
gizing it  (pp.  186-195). 

This  experiment  proves  beyond  question  that  the  throat- 
apparatus  in  the  tortoise  is  essential  to  life,  while  the  greater 
develoj)ment  than  in  the  frog  is  due  to  the  fact  of  the  modifi- 
cations in  the  body-envelope  inhibiting  the  downward  move- 
ment in  the  floor  of  support  to  the  viscera  for  facilitating  lung- 
diastole,  the  organs  being  compelled  to  act  upon  a  horizontal 
plane  in  a  to-and-fro  movement  upon  the  viscera.  At  the  same 
time,  however,  it  involves  independent  automatic  action  in  the 
lungs,  for  in  no  other  way  could  the  air  be  effectively  diffused 
through  the  alveolar  compartments  in  the  measure  of  the  phy- 

*  This  was  done  in  London,  that  would  not  suffer  physiological  investiga- 
tions, its  heart  too  tender.  All  the  same,  I  made  it  out  there  ;  this  and  frog- 
respiration,  with  other  important  matters,  all-pervading  Intelligence  blessing 
the  time  and  place. 


420 


EFFECT   UPON   CAKDIAC   DEVELOPMENT. 


siological  requirements.  Cut  the  point  we  wish  to  make  in  this 
connection,  is  the  effect  produced  by  this  embarrassed  respira- 
tion upon  cardiac  development,  since  it  throws  more  labor  upon 
the  heart.  This  circumstance  is  seen  in  the  great  increase  in 
the  muscles  of  the  heart  (Fig.  175,  A'),  the  walls  of  the  ven- 


Fig.  175. — The  Viscera  in  the  Tortoise  (Emys  Europced),  as  viewed  from  below. — 
Bojamis.  A',  heart  (ventricle)  ;  A,  aorta  ;  B',  right  auricle  ;  B",  left  auricle  ;  B, 
left  aorta  ;  C,  common  pulmonary  artery  ;  D,  right  subclavian  artery  ;  E,  trachea  ; 
E\  left  ;  E",  right  carotid  arteries  ;  i,  jugular  ;  h,  brachial  veins  ;  e,  g,  vertebral 
and  subclavian  veins  ;  T,  T,  oesophagus  ;  K,  stomach,  nearly  covered  by  the  liver  ; 
K',  pylorus  ;  7,  J,  liver  ;  Z,  Z,  lungs  ;  31,  oviducts  ;  iV,  ovaries  ;  V,  intestine  ;  £7, 
urinary  bladder  ;  U',  U",  anal  bursse  ;  X,  cloaca  opening. 

tricle  being  much  thicker  comparatively  than  in  other  animals 
(Figs.  176,  ]  77).  That  this,  together  with  the  sluggish  movements 
of  the  animal,  is  due  to  defective  action  in  the  lungs,  has  forci- 
ble proof  in  the  case  of  the  flying  lizard  (Fig.  179),  in  which 


EFFECT  UPON  CAEDIAC  DEVELOPMENT. 


427 


respiration  is  free,  the  movements  energetic,  the  heart  (a)  com- 
paratively small. 

The  reduced  size  and  number  of  the  muscles  in  the  ventricle 
are  coincident  with  the  increased  and  freer  respiration,  while 
this  provides  for  corresponding  increase  in  the  activities,  which 
must  have  their  equivalent  in  force  evolved  by  means  of  respi- 
ration, the  two  being  in  correspondence,  from  the  very  na- 
ture of  things.  At  the  same  time,  this  indicates  a  struggle 
with  some  force,   which  is  gravitation  ;  hence  the  extensive 

Fig.  176.  Fig.  177. 


Heart  of  Tortoise  (Chelys  fimbriate). — Owen.  Showing  interior  of  ventricle,  the  incom- 
plete interventricular  septum  dividing  the  aortic  from  the  pulmonic  cavities,  through 
which  the  bristle  is  passed  in  the  two  sections. 

Fig.  176. — P,  pulmonary  artery  laid  open  at  the  root,  so  as  to  expose  the  bivalved  orifice, 
the  bristle  passed  through  the  incomplete  septum. 

Fig.  177.  —A',  right  aorta  ;  A,  left  aorta,  similarly  exposed,  and  the  incomplete  septum 
cut  through,  the  bristle  passed  through  the  pulmonary  vein  of  that  side  ;  the  root  of 
right  aorta  (A')  is  behind  that  of  the  left  ;  the  elongated  rounded  bodies  are  the 
auricles. 

provisions  in  land  animals  for  generating  force  in  the  organ- 
ism, the  increase  in  respiratory  surface,  greater  development 
of  the  vascular  system,  and  the  numbers  of  the  muscles  and 
nerves  to  the  heart  and  blood-vessels  for  effecting  rapid  impor- 
tation and  distribution  of  the  force-producing  elements,  together 
with  the  rise  and  fall  in  frequency  of  the  respiratory  and  cardo- 
arterial  rhythms  with  the  swell  in  the  activities,  the  increased 
respiration  necessarily  involving  a  corresponding  increase  in 
circulation  in  order  to  make  it  effective,  the  objective  point 
being  the  cell-brood  in  the  tissues.  In  lizards  (Fig.  179),  in 
which  the  voluntary  movements  are  rapidly  performed,  the 
lungs  are  more  highly  developed,  reduced  in  size,  and  more 
compact  from  multiplication  of  the  septa  and  capillary  plex- 


426  THE   INTEKVENTRICULAK    SEPTUM. 

uses  ;  and  by  reason  of  the  differentiation  of  "  ribs  "  for  assist- 
ing and  facilitating  expansion  and  contraction  in  the  lungs  and 
body-walls,  freer  respiration  is  provided  for,  whereby  more 
energetic  actions  are  enabled  than  are  possible  to  chelonia,  the 
effect  at  the  same  time  being  to  relieve  the  heart  in  proportion, 
obviating  the  necessity  for  the  number  of  the  muscles  that  ob- 


Fig.  178. — Structure  of  Auricles  ;  Heart  of  Chehjs  fimbriata.—  Owen.  The  left  auricle 
(M)  receives  the  arterial  blood  from  the  lungs  by  a  single  vein,  the  common  trunk 
of  the  pulmonary  veins  (e,  e)  ;  it  opens  into  the  back  part  of  the  auricle,  near  the 
septum,  and  is  guarded  by  a  single  oblique  membranous  fold  (J/).  Each  auricular 
ventricular  orifice  is  guarded  by  a  fold  which  extends  across  it  from  either  side  of 
the  base  of  the  inter-auricular  septum  ;  to  that  of  the  left  auricle  a  small  part  of  the 
muscular  structure  is  attached  by  chordae  tendinae.  Opposite  to  the  right  valve  a 
semilunar  ridge  projects,  in  Tcstudo  indica,  whLh  is  the  rudiment  of  the  second 
auriculo- ventricular  valve  in  the  Crocodile,  and  of  the  fleshy  valve  of  that  orifice  in 
the  right  ventricle  of  Birds.  The  apex  of  the  ventricle  is  attached  by  a  short  fold  of 
the  serous  membrane  to  the  pericardium  (t).  The  bivalved  orifice  between  the  sinus 
and  the  auricle  (i\  o)  is  a  transverse  slit.  The  white  arrow  (o)  shows  the  course  of 
the  blood  from  the  right  auricle,  past  the  valve  supported  by  the  base  of  the  auric- 
ular septum,  into  the  aortic  cavity  of  the  ventricle.  In  Fig.  176,  a  bristle  passes 
through  the  orifice  left  by  the  incomplete  septum  between  the  aortic  and  pulmonary 
cavities  into  the  latter,  which  is  largest  in  Emys,  as  in  Ophidia  and  Lacertilia;  the 
bivalved  orifice  of  the  pulmonary  artery  (P)  is  shown.  In  Fig.  177,  that  of  the  left 
aorta  (A)  is  similarly  exposed,  and  the  incomplete  septum  is  cut  through  ;  the  root 
of  the  right  aorta  (-4.0  is  behind  that  of  the  left.  The  relative  position  of  the  origins 
of  the  three  arteries  from  the  chelonian  ventricle  is  shown  in  Fig.  177,  where  (P)  is 
the  pulmonary,  (-4)  the  left  aorta,  (-4')  the  right  aorta — the  most  posterior,  or  dorsal, 
of  the  three  arteries. 

tain  in  the  heart  of  chelonia.  Then,  again,  in  birds,  in  which 
the  voluntary  actions  are  enormously  increased  in  force,  we 
find  here  also  that  respiration  and  circulation  are  in  corre- 
spondence ;  otherwise,  they  could  not  be  produced.  It  follows, 
with  progress  in  development  in  which  more  and  more  force  is 
called  for,  that  commensurate  arrangements  must  obtain  for 


HEAET  AND   LUNGS. 


429 


arterializing  and  circulating  the  blood  ;  hence  the  differentia- 
tion of  the  left  cardiac  chambers  for  separating  the  pulmonic 
or  aerated  blood  from  the  systemic  venous  current,  the  limi- 
tation of  the  arterial  or  aortal  trunk  to  the  left  side  of 
the  heart,  the  increase  in  respiratory  surface,  together  with. 


Fig.  179. — Abdominal  Cavity  and  Viscera,  Draco  Volans. — Cyclopediae  of  Anatomy  ; 
Art.  Reptilia.  By  T.  Rymer  Jones,  a,  Ventricle  of  heart  ;  b,  right  auricle  ;  c,  left 
auricle  ;  d,  carotid  artery  ;  e,  jugular  vein  ;  /,  subclavian  artery  ;  g,  trachea  ;  ky 
liver  ;  I,  lower  venous  sinus,  commencing  from  liver  and  extending  to  right  venous 
sinus  ;  m,  biliary  duct  ;  o,  stomach  ;  p,  beginning  of  small  intestine  ;  q,  small  intes- 
tine ;  r,  beginning  of  large  intestine  ;  s,  its  attenuated  region  ;  t,  t,  kidneys  ;  u,, 
cloaca  ;  v,  bladder  ;  w,  anus. 

the  high  pressure  which  obtains  in  the  arterial  system  for  in- 
creasing circulation  in  the  capillaries  and  cell-brood,  which  is 
gradually  brought  about  in  the  reptilia  and  perfected  with  the- 
birds.  .\nd  the  crocodile  is  the  connecting  link  between  the 
birds  and  reptiles,  the  interventricular  septum  being  perfected 


430  CROCODILE   CONNECTING   LINK    WITH  THE   BIRDS. 

in  the  crocodile,  but  leaving  an  opening  between  the  two  aortse, 
causing  the  venous  blood  to  flow  through  the  body  during  sub- 
mergence, when,  by  reason  of  defective  respiration,  the  blood 
tends  to  accumulate  at  the  right  side,  and  but  for  these  safety- 
valves,  life  would  be  impossible  ;  at  the  same  time,  it  should 
promote  the  nutritive  processes,  for  the  reasons  already  given 
(pp.  366-369).  The  anatomical  peculiarities  in  the  heart  of  the 
crocodile  are  succinctly  stated  in  the  following  excerpt : 

' '  In  the  Crocodilian  order  a  marked  advance  is  made  in  the 
structure  of  the  heart.  The  blood  from  the  general  system  is 
poured  by  the  veins  into  a  sinus  (Fig.  180,  S),  whence  it  passes 
into  a  right  auricle  (o)  by  the  usual  bivalved  aperture.  The 
auricle  has  a  more  distinct  'appendix,'  and  its  muscular  walls 
are  thicker  than  in  lower  reptiles.  The  auriculo-ventricular 
orifice  is  defended  not  only  by  the  ordinary  valve  on  its  left 
side,  which  is  attached  to  the  base  of  the  auricular  septum,  but 
by  a  similar  though  smaller  fold  on  the  opposite,  or  right  side ; 
this  fold  becomes  the  fleshy  auriculo-ventricular  valve  in  birds. 
To  the  junction  of  the  two  valves  at  their  lower  angle  a  fleshy 
column  is  attached.  The  ventricular  cavity  (i«J),  which  receives 
the  venous  blood,  propels  it  to  the  left  aorta  (A)  and  to  the 
pulmonary  artery  (P) ;  the  origin  of  each  is  guarded  by  a  pair 
of  semilunar  valves.  Immediately  above  the  larger  of  those 
of  the  left  aorta  is  an  orifice  leading  into  the  right  aorta ;  in 
Fig.  180,*  a  bristle  is  passed  from  the  left  aorta  through 
this  orifice  into  the  right  axillary  branch  (a)  of  the  right  or 
brachio-cephalic  aorta.  In  the  figure,  the  valve  is  drawn  down 
to  show  the  orifice  ;  in  its  natural  state  it  conceals  and  would 
-cover  the  orifice,  as  the  blood  flowed  from  the  ventricle  into  the 
left  aorta.  Some  openings  lead  from  the  pulmonic  cavity  of 
the  ventricle  into  a  spongy  structure,  which  has  been  defined 
as  a  particular  cavity  {spatium  inter •ventricular e)  of  the  ven- 
tricle ;  but  it  is  essentially  a  part  of  the  pulmonic  chamber ; 
bristles  are  passed  through  the  orifices  or  intercolumnar  spaces 
leading  from  M  to  this  structure.  The  left  auricle  (Fig.  18,  1, 
M\),  when  distended,  is  smaller  than  the  right,  and  of  a  more 
transverse  form  ;  its  muscular  part  is  produced  into  an  append- 
age, which  almost  meets  that  of  the  right  auricle  in  front  of 

*  Ibid,  339.  t  Ibid,  340. 


THE   CROCODILIAN    HEART. 


431 


the  '  conus  arteriosus,'  embracing  the  'sulcus  coronalis'  of 
the  heart.  There  is  a  small  pulmonary  sinus  receiving  the 
short  trunks  of  the  pulmonary  veins  (I,  T).  The  left  auriculo- 
ventricular  aperture  is  defended  by  a  broad  membranous  fold 
continued  into  the  ventricle  from  the  middle  of  the  base  of 
the  interauricular  septum ;  to  its  margin  are  attached  a  few 


Fig.  180. — Right  Auricle  and  Ventricle,  Crocodilus  Acutiis. — Owen.  P,  P,  right  and 
left  pulmonary  arteries  laid  open  at  the  root,  so  as  to  expose  the  semilunar  valves 
(P),  a  bristle  passed  thence  through  right  pulmonary  artery  :  Ah,  left  aorta  ;  A', 
right  aorta  ;  A",  intercommunicating  channel  ;  rx,  superior  vena  cava  ;  v,  inferior 
vena  cava  ;  s,  sinus  ;  o,  right  auricle  ;  R,  right  ventricular  cavity  ;  ax,  as,  right 
and  left  carotid  arteries  ;  a,  a,  right  and  left  subclavian. 

chordse  tendinese ;  the  cavity  into  which  it  opens  {L)  is  dis- 
tinct from  the  pulmonic  cavity,  the  septum  being  complete  ;  its 
walls  are  smooth,  or  less  broken  by  columnse  cornese  than  in 
other  reptiles ;  and  the  free  walls  of  this  ventricle  are  more 
compactly  muscular.  The  ventricle  is  produced  in  a  sub- 
conical  form,  from  its  base  to  the  origin  of  the  right  or  brachio- 
cephalic aorta  ;  the  auriculor  ventricular  valve  is  slit,  to  show 
the  course  of  the  ventricle  to  the  origin  of  that  aorta  ;  this  has 


432  THE   CKOCODILIAN   HEAET. 

a  pair  of  semilunar  valves,  above  which  is  the  intercommuni- 
cating orifice  with  the  left  aorta.  Thus,  the  heart  in  Crocodilia 
consists  of  two  auricles  and  two  ventricles,  corresponding  to 
the  'right'  and  'left'  auricles  and  ventricles  of  mammals. 
But,  through  the  origin  of  an  aorta  from  the  right  as  well  as 
from  the  left  ventricle,  and  their  intercommunication,  it  fol- 
lows that  whenever,  from  an  impeded  state  of  the  pulmonary 
circulation,  the  right  ventricle  and  its  arteries  become  over  dis- 
tended, the  venous  blood  flows  through  the  inter-aortic  orifice 
into  the  arterial  trunk,  which,  after  supplying  the  head  and 
f orelimbs,  bends,  at  A' ,  over  the  right  bronchus,  and  effects  a 
union  at  A"  (Fig.  lbO)  with  the  left  aorta,  A,  h.  Such  a  state 
of  the  circulation  coincides  with  and  facilitates  the  long  sub- 
mersion of  the  crocodile.  When  the  animal  is  on  land,  and 
breathing  the  air  directly,  the  arterialized  blood  flows  freely 
into  the  ventricle  (lbl,  L),  and  the  synchronous  currents  from 
this  and  the  opposite  ventricle  throw  forward  the  valves  at  the 
respective  origins  of  the  two  aortse  and  close  the  inter-aortal 
orifice.  The  arterial  and  venous  streams  flow  on  unmixed : 
the  former  to  the  brain  and  other  parts  of  the  head  and  fore- 
limbs  ;  the  latter,  by  the  branch,  7i  (Fig.  180),  chiefly  to  the 
liver  and  contiguous  viscera ;  a  small  part  mixing  with  the 
arterial  blood  in  A',  to  be  transmitted  by  A"  to  the  other 
abdominal  viscera,  hind  limbs  and  tail.  To  convert  the  heart 
of  the  crocodile  into  that  of  the  bird,  it  needs  only  to  obliterate 
the  left  aorta,  to  appropriate  the  right  or  pulmonic  ventricle 
(Fig.  ISO,  JR)  exclusively  to  the  service  of  the  pulmonary 
artery  ;  and  the  '  left '  or  systemic  ventricle  to  the  service  of 
the  aorta,  which,  in  Hazmototherma  (warm-blooded  animals)  is 
the  exclusive  distributer  of  arterial  blood,  in  an  unmixed  state, 
to  the  general  system." 

Coming  to  the  birds,  keeping  in  mind  the  cardinal  circum- 
stance in  evolution,  or  the  struggle  with  gravitation,  together 
with  the  means  for  evolving  force  in  the  organism  or  by  respi- 
ration, inclusive  of  the  actions  in  the  heart  and  vessels,  since 
the  whole  f onus  a  connected  movement,  as  before  remarked,  we 
have  only  to  regard  the  mechanics  from  that  stand-point,  and 
the  whole  is  at  once  made  intelligible.  Thus,  the  animal  is 
lifted  off  the  ground  in  place  of  lying  at  full  length  upon  it, 


PKINCTPLE   IN  CAEDIAC   DEVELOPMENT. 


483 


and  supported  by  two  chains  of  bones  forming  the  cn.ra, 
which  compel  eternal  vigilance  and  a  tremendous  struggle 
with  gravitation  in  order  to  effect  it,  the  animal  literally  living 
upon  its  feet,  the  body  at  greater  or  less  elevation  off  the 
ground,  tending  to  fall  to  the  earth  all  the  time,  and  with 
every  movement  to  topple  over  in  the  direction  of  propelling 


i 

Pig.  181.—  Left  Auricle  and  Ventricle,  Crocodilus  Acutus. — Owen.  M,  left  auricular 
cavity  ;  L,  left  ventricular  cavity  ;  V**,  inferior  vena  cava  ;  S,  sinus  formed  at  the 
junction  of  upper  and  lower  cava  ;  I,  I,  short  trunks  of  pulmonary  veins  ;  P,  P , 
pulmonary  arteries  ;  D,  Df,  D',  intercommunicating  channel  between  pulmonary 
arteries  and  aortse  ;  ax,  a*,  carotid  arteries  ;  a,  a,  subclavian  arteries  ;  5,  6,  the 
two  innominata.    The  pulmonary  arteries,  veins  and  upper  cava  are  ligated. 

force.  Marvelous  adjustments  with  gravitation  obtain  in  it, 
but  for  obvious  reasons  we  shall  at  present  only  consider  the 
effect  upon  cardiac  development. 

For  evolving  the  requisite  amount  of  force  in  the  bird,  most 
extensive  respiratory  arrangements  obtain  in  the  lungs  (Fig. 
182)  and  air-bladders,  with  which  the  lungs  communicate  by 
means  of  openings  (a,  e)  carried  through  the  lung-substance, 


434  PRINCIPLE   IX  CAKDIAC   DEVELOPMENT. 

the  air  penetrating  the  very  bones  themselves  by  means  of  an 
extensive  canalicular  system,  and  in  birds  of  vigorous  flight 
passing  into  sacs  between  the  muscles  of  the  chest  and  neck  ; 
while  for  increasing  the  action  during  flight  a  special  arrange- 
ment obtains  in  the  costal  framework  by  the  formation  of 
elbows  (Fig.  23),  by  means  of  cartilages  in  the  mid-costal  re- 
gion, dividing  the  ribs  into  "superior"  and  "inferior"  (/,  j), 
whereby  the  action  in  the  chest  is  greatly  facilitated,  at  the 
same  time  firmness  and  strength  are  imparted  to  the  dorsal 
portions ;  while  by  means  of  the  muscles  of  the  abdomen, 
which  are  inserted  into  them  and  the  sternum,  the  chest  is 
rhythmically  expanded  and  contracted  by  the  action  in 
the  abdomen  during  respiration ;  while  during  flight  the 
chest  is  rhythmically  expanded  and  contracted  by  the  alter- 
nate extension  and  retraction  of  the  wings  (p.  79),  so  that 
respiration  is  necessarily  compelled  to  be  in  correspondence 
with  the  activities  during  the  time  of  greatest  exertion,  in 
order  to  evolve  the  additional  amount  of  force  thereby  ex- 
pended. Such,  in  brief,  is  the  ample  provision  for  generating 
force,  but  which  calls  for  corresponding  provision  in  the  circu- 
latory organs  in  order  to  make  it  eifective.  Accordingly,  we 
have  the  heart  and  blood-vessels  completely  developed  ;  at 
the  same  time,  the  number  of  the  muscles  and  nerves  are 
increased  correspondingly  in  both,  while  in  the  arterial  sys- 
tem we  have  high  blood-pressure  for  speeding  the  blood  in 
the  capillaries  for  energizing  the  local  actions,  while  this,  in 
turn,  calls  for  the  thick,  strong  elastic  coat  in  the  arteries  in 
order  to  produce  this  increased  pressure  in  the  arterial  sys- 
tem ;  while  the  whole  relates  to  the  generation  of  force  in 
the  organism  commensurate  with  the  amount  expended  in 
producing  the  activities. 

Say  what  you  will  of  life  !  The  evolution  of  force  in  the 
organism  by  means  of  respiration  and  circulation,  which  are 
based  upon  rhythmical  changes  in  pressure,  invoked  in  the 
measure  of  the  requirements,  is  the  law  regulating  the  vital 
phenomena,  anatomical  and  physiological,  in  animal  life.  It 
is  needless  to  extend  the  matter. 

The  following  illustration  (Fig.  1S3)  exhibits  the  mode  of 
termination  of  the  great  venous  trunks  at  the  heart  in  reptiles 


PRINCIPLE   IN  CARDIAC   DEVELOPMENT. 


435 


and  warm-blooded  animals,  and  the  changes  with  progress  in 
development. 


Pig.  182.— Heart  and  Great  Vessels  of  Buteo  Vulgaris.  — Gegenbaur.  tr,  Trachea ;  tr 
crop  ;  ae,  communication  between  the  air-sacs  and  the  lungs  ;  b,  Bursa  Fabricii  ;  aor 
aortic  arch  ;  aad,  right  anonymous  artery  ;  acts,  left  anonymous  artery  ;  ps,  left 
pulmonary  artery  ;  c,  carotid  ;  am,  visceral  artery  ;  vci,  commencement  of  the 
inferior  vena  cava  ;  vcm,  artery  to  coccygeal  and  mesenteric  regions. 

In  Fig.  184,  we  have  the  different  stages  in  development  of 
the  proximal  arterial  vessels  in  embryological  evolntion,  to  be 


430 


EVOLUTION    OF   AKTEEIAL   VESSELS. 


read  from  left  to  right,  beginning  by  the  formation  of  the 
truncus  arteriosus  with  one  vascular  loop  only,  the  ones  imme- 
diately succeeding  this  being  shaded  in,  then  the  upper  ones  ; 
making  five  pairs  of  vascular  loops  in  all,  as  in  the  fishes  (2);  fol- 

t  I  31  iv 


Eijr.  183.— Showing  Mode  of  Termination  of  the  Great  Venous  Trunks  in  Different 
Stages  in  Development,  viewed  from  behind. — Gegenbaur.  I,  Reptile  (Python)  ;  II, 
Bird  (Larcorhamphus)  ;  III,  Marsupial  (Halmaturus)  ;  IV,  Pig  ;  i,  Vena  cava  inferior ; 
s,  vena  cava  superior  sinistra  ;  d,  vena  cava  superior  dextra  ;  ap,  pulmonary  artery ; 
si;  sinus  venosus. 


Pig.  1S4. — Showing  the  Different  Stages  in  Development  of  the  Proximal  Arterial 
Vessels  in  Embryological  Evolution.—  Carpenter.  1,  Truncus  arteriosus  with  one 
pair  of  aortic  arches,  and  dotted  outlines  indicating  the  future  position  of  the  second 
and  third  pairs  ;  2,  truncus  arteriosus  with  four  pairs  of  aortic  arches  and  indica- 
tions of  the  fifth  ;  3,  truncus  arteriosus  with  the  three  posterior  pairs  of  aortic 
arches,  from  which  the  permanent  vessels  of  the  embryo  are  developed,  with  dotted 
outlines  showing  the  position  of  the  two  (now)  obliterated  anterior  arches  ;  4,  per- 
manent arterial  trunks  in  their  primitive  form,  the  obliterated  portions  still  shown 
in  dotted  outline  ;  1-5,  primitive  aortic  arches  ;  ta,  truncus  arteriosus  ;  a,  aorta  ; 
p,  pulmonary  artery  ;  p',  p",  branches  to  the  lungs  ;  aw',  root  of  thoracic  aorta  (ad) 
on  left  side  ;  aw,  obliterated  root  springing  from  right  side  ;  s',  s",  subclavian 
artery  ;  v,  vertebral ;  ax,  axillary  ;  c,  common  carotid  ;  c',  external  carotid  ;  c", 
internal  carotid. 

lowed  by  the  gradual  obliteration  which  sets  in  after  a  certain 
stage  in  development  is  reached  ;  till,  finally,  we  have  the  con- 
dition in  4,  representing  the  permanent  vessels  or  aorta,  pul- 
monary, carotid,  subclavian  and  vertebral  arteries. 


EVOLUTION    OF   ARTERIAL   VESSELS.  437 

In  the  first  two  pairs,  or  highest  arches,  certain  portions  are 
obliterated,  leaving  three  pairs  ;  of  these,  the  lowest  pair  is  en- 
tirely obliterated  on  the  right  side  ;  on  the  left,  it  gives  off  the 
pulmonary  artery,  and  remains  throughout  foetal  life  the  chan- 
nel of  communication  with  the  aorta,  or  ductus  arteriosus; 
the  bulbus  aorticus  is  subdivided  by  adhesion  of  its  walls,  there- 
by forming  two  tubes,  one  of  which  is  the  nascent  aorta,  the 
other  pulmonary  artery ;  while  of  the  second  pairs,  the  right 
arch  forms  the  innominate  and  first  portion  of  right  subclavian ; 
the  left  becomes  the  arch  of  the  aorta,  and  contributes  to  form 
the  left  subclavian.  The  third  pair  contributes,  with  portions 
of  the  two  highest  pairs,  to  the  formation  of  the  internal  and 
external  carotid  arteries. 

The  following  beautiful  illustration  (Fig.  185),  by  an  English 
naturalist, f  affords  a  bird's-eye  view  of  the  vascular  changes 
which  accompany  the  transition  from  intra-uterine  to  extra- 
uterine life,  while  the  mechanical  principle  which  underlies 
these  changes  in  the  mechanics  of  circulation  is  fully  set 
forth  in  the  chapter  upon  the  embryonic  circulation  (pp.  338, 

341). 

Before  leaving  this  subject,  I  desire  to  call  attention  to  a 
deeply  suggestive  fact  connected  with  circulation  in  the  foetus, 
notably,  the  very  small  amount  comparatively  of  arterialized 
blood  which  actually  reaches  the  heart  of  the  embryo,  which 
may  be  roughly  estimated  at  one-fourth  only  of  the  blood  in 
the  umbilical  vein,  the  great  bulk  of  the  blood  passing  through 
the  liver  parenchyma  for  effecting  assimilation  and  promoting 
Tisematosis  ;  tlae  liver,  undoubtedly,  being  concerned  in  both, 
is  also  relatively  larger  than  at  any  other  period  of  life  ;  and 
since  this  would  necessarily  consume  the  oxygen,  it  follows 
that  the  supply  would  have  to  be  by  the  route  of  the  ductus 
venosus  (7);  while  the  small  size  of  the  canal  is  of  itself  eloquent 

f  "  Outlines  of  Comparative  Anatomy  :  Presenting  a  sketch  of  the  present 
state  of  knowledge,  and  of  the  progress  of  discovery,  in  that  science ;  and 
designed  to  serve  as  an  introduction  to  animal  physiology ,  and  to  the  principles 
of  classification  in  zoology."  By  Robert  E.  Grant,  M.D.,  F.R.S.,  Lond.  &Ed., 
F.L.S.,  F.G.S.,  F.Z.S.,  M.W.S.,  etc. ;  Fellow  of  the  Royal  College  of  Physicians 
of  Edinburgh  ;  late  Professor  of  Physiology  in  the  Royal  Institution  of  Great 
Britain,  and  Professor  of  Comparative  Anatomy  and  Zoology  in  University 
College,  London.     1841. 


4:38 


VASCULAR  CHANGES   AT   BIRTH. 


m- 


Fig.  185. — Circulation  in  Intra  and  Extra  Uterine  Life,  showing  the  vascular  changes. — 
Grant.  B,  intra  uterine  circulation  in  the  matured  embryo  ;  A,  extrauterine  B,  4, 
umbilical  vein  ;  5,  sinus  formed  by  junction  of  umbilical  vein  with  vena  portae  ; 
1,  2,  3,  vena  portae  ;  G,  hepatic  veins  (principally  right  lobe,  unmarked  vessel  indi- 
cating left  lobe  of  liver)  ;  7,  ductus  venosus  ;  10,  1 1 ,  inferior  vena  cava  ;  cZ,  right 
auricle  ;  e,  right  ventricle  ;  g,  ductus  arteriosus  ;  k,  junction  with  first  portion  of 
descending  aorta,  pulmonary  artery  dividing  into  right  and  left  pulmonary  arteries  (h, 
i)  beyond  ductus  arteriosus  ;  /,  left  auricle  ;  I,  ascending  arch  of  aorta;  m,  arteria 
innominata  giving  off  carotid  and  subclavian  of  right  side  ;  n,  left  carotid  ;  o,  left 
subclavian  artery  ,  s,  right  common  iliac  ;  r,  left  common  iliac  artery  ;  u,  v,  um- 
bilical arteries,  continuation  of  internal  iliac  ;  t ,  right  external  iliac  artery  ;  iv,  left 
external  iliac  ;  8,  9,  right  and  left  external  iliac  veins  ;  a,  b,  brachio-cephalic 
trunks  ;  c,  superior  vena  cava  ;  h,  right  pulmonary  arter}-  ;  i,  left  pulmonary  ar 
tery.  A,  I,  pulmonary  artsiy ;  m,  left  branch;  n,  right  branch;  p,  atrophied  remains 
of  ductus  arteriosus  ;  /,  left  auricle  ;  g,  ascending  aorta  ;  h,  right  carotid  from 
innominata  ;  i,  left  carotid  ;  k,  left  subclavian  artery  ;  r,  s,  diaphragmatic,  cceliac, 
and  superior  mesenteric  arteries  ;  t,  inferior  mesenteric  artery  ;  u,  spermatic  ar- 
tery ;  iv,  r,  common  iliac  arteries  ;  e,  x,  external  iliac  ;  y,  y,  internal  iliac  arteries ; 
1,  1,  external  iliac  ;  2,  2,  internal  iliac  veins  ;  3,  3,  common  iliac  veins  ;  4,  inferior 
vena  cava  ;  5,  right  spermatic  vein  ;  6,  inferior  mesenteric  vein  :  7,  right  renal 
vein  ;  10,  11,  12,  venae  portae  ;  13,  14,  divisions  of  same  which  ramify  in  right  and 
left  lobes  of  the  liver  ;  8,  9,  terminations  of  the  hepatic  veins  in  the  vena  cava  (the 
portal  vessels  being  moved  down  in  order  to  show  this  circumstance). 


LITTLE    OXYGEN   NEEDED   IN   NUTRITION.  439 

proof  that  but  a  relatively  small  proportion  of  oxygen  is  con- 
sumed in  embryonic  evolution.  Now,  then,  for  the  question: 
Why  should  the  embryo  call  for  the  minimum  of  arterial 
blood?  This  also  is  readily  answered,  notably :  1.  Oxygen  is 
the  force-producer  for  evolving  heat  and  producing  motion  in 
animal  organisms  ;  but  since  the  maternal  tissues  are  sufficient 
for  maintaining  temperature,  temperature  in  the  abdomen  be- 
ing the  highest  in  the  body,  the  embryo  would  not  need  oxy- 
gen for  this  purpose  any  more  than  the  chick  in  incubation. 
2.  The  nutritive  processes  per  se  do  not  require  much  oxygen, 
the  amount  consumed  in  the  liver  being  simply  by  way  of 
preparation,  so  to  speak,  in  blood-making,  the  production  of 
cells  and  albuminoid  compounds,  etc.  ;  while  the  crystalliza- 
tions in  tissue-structure,  as  is  the  case  in  plants,  would  call 
for  but  slight  expenditures  in  order  to  effect  them  ;  in  prooZ 
of  which,  we  have  only  to  refer  to  circulation  below  the  dia- 
phragm. 

The  structures  below  the  diaphragm  in  actual  weight  exceed 
those  above  it,  and  by  universal  admission  no  arterial  blood  is 
ever  sent  into  this  portion ;  nevertheless,  nutrition  is  very 
active.  Furthermore,  when  you  take  into  consideration  the 
hepatic  circulation  and  the  small  amount  of  arterial  blood 
reaching  the  lower  cava  ;  that  this  is  first  sent  to  the  brain 
and  upper  portions  of  the  body  before  reaching  the  heart  again 
through  the  upper  cava,  thence  through  the  ductus  arteriosus 
to  the  lower  portions,  that  in  the  very  nature  of  things  it  can- 
not contain  much  oxygen  ;  on  the  contrary,  is  highly  venous. 

Facts  are  stubborn  things ;  none  more  so  than  anatomy. 
And  who  would  read  them  aright,  must  regard  them  from  the 
stand-point  furnished  in  the  organic  laws  and  the  history  in 
development.  It  comes  to  this,  then,  that  nutrition  may  be 
very  active  without  much  oxygen,  the  importance  of  which  it 
would  be  difficult  to  overestimate,  since  it  would  establish 
kindred  processes  in  the  nutrition  of  animals  and  plants, 
for  which  we  have  been  contending.  Look  into  it  well,  please  ; 
it  is  no  hasty  generalization,  based  upon  insufficient  data,  but 
a  sober  fact,  proven  upon  the  anatomy  of  the  foetus,  there- 
fore incontrovertible. 

3.  But  for  producing  motion,   this  would  call  for  oxygen 


440  occasion  fo::  a  ductus  venoscs. 

sufficient  only  to  keep  up  the  action  in  the  cardiac  and  vas- 
cular centres,  as  respiration  proper  does  not  go  on,  and  there  is 
no  occasion  for  expenditure  of  force,  save  in  the  heart  and 
blood-vessels  ;  and  should  any  other  movements  occur,  notably 
the  reflex  actions  in  the  spinal  cord,  as  when  the  foetus  is  said 
"  to  be  active,"  we  may  depend  upon  it  that  the  ductus  arteri- 
osus is  unusually  large.  It  is  a  mistake  to  fancy  that  the  brain 
in  seds  oxygen  for  effecting  growth;  the  facts  established  by 
daily  experience  prove  the  contrary,  notably  the  phenomena  in 
sleep,  especially  in  cases  in  which  sleep  is  induced  artificially 
by  full  doses  of  morphia  and  hydrate  of  chloral,  in  which  the 
respirations  are  nearly  suspended,  the  patient  coming  out  of 
this  condition,  nevertheless,  vastly  improved  (p.  367).  Yes,  it 
is  an  error  to  refer  the  nutritive  processes  in  the  brain  or  any 
other  tissue  to  the  presence  of  oxygen  in  the  blood.  On  the 
contrary,  everything  goes  to  establish  that  the  animal  tissues,  in 
common  with  vegetal,  are  best  nourished  in  fluids  saturated 
with  carbonic  acid  gas,  and  there  is  no  controverting  that  fact. 
Why  seek  to  controvert  it?  IS  ay,  keep  this  in  mind  when 
you  do  so  :  Force  one  thing,  nutrition  another,  and  quite  the 
opposite. 

The  former  effects  re-distribution,  disintegrates  and  dis- 
perses matter;  the  latter  masses  it,  brings  it  together  in 
methodical  order,  effecting  increase  in  size.  Oxygen  is  the 
agent  for  effecting  the  one,  carbonic  acid  the  other,  at  least  is 
the  principal  agent  in  plants  ;  while  nitrogen  is  added  in  greater 
or  less  proportion  in  all,  but  greatest  in  animal  tissue. 

The  head  downward  (which  is  the  usual  position),  with  the 
force  in  the  umbilical  vein  and  the  suction-force  in  the  heart, 
the  blood  from  the  ductus  venosus,  liver  and  lower  cava  are 
readily  drawn  into  the  heart,  passing  through  the  right  to  the 
left  auricle  and  ventricle,  guided  by  the  Eustachian  valve,  and 
undergoing  admixture  along  the  route  and  in  the  ventricle,  is 
sent  thence  to  the  cardiac  structures  and  brain,  the  small 
amount  of  oxygen  it  contains  being  sufficient  for  evolving  the 
requisite  force  in  the  cardiac  and  vaso-motor  centres,  which 
are  the  principal  ones  in  habitual  action. 

The  coronary  arteries  springing  from  the  root  of  the  aorta, 
immediately  above  the  valves,  insure  prompt  supply  to  the 


OCCASION"   FOR  A  D    CTUS   VENOSUS.  44L 

cardiac  muscles  and  nervous  ganglia,  while  the  carotid  and 
vertebral  arteries  maintain  the  activities  in  the  vaso-motor  cen- 
tre, so  that  nervous  force  is  readily  evolved  for  producing  the 
pumping  actions  in  the  heart  and  vessels,  making  this  com- 
mensurate with  the  nutritive  processes.  All  which  is  plain 
enough  from  the  stand-point  furnished  in  the  organic  laws 
underlying  the  organism  ;  otherwise,  are  utterly  inexplicable. 
And  with  the  great  respiratory  movement  in  abeyance,  the 
body-temperature  maintained  by  the  maternal  tissues,  volition 
and  the  voluntary  movements  wrapped  in  profoundest  slumber 
and  made  impossible  from  the  absence  of  the  force-producer, 
oxygen,  the  action  in  the  heart  and  vessels  only,  inclusive,  of 
course,  of  the  action  in  the  cell-brood,  the  nutritive  processes 
are  more  closely  approximated  to  the  floral  than  at  any  other 
period.  The  principle  in  nutrition  is  in  the  ascendency,  while 
the  one  for  evolving  force  by  respiration  must  wait  till  all  things 
are  ready  and  the  change  in  environment  comes.  How  small 
the  ductus  arteriosus  (Fig.  135,  7)  contrasted  with  the  size  of 
the  umbilical  (4)  and  hepatic  veins  !  (!>) — but  taking  the  shortest 
and  most  direct  route  to  reach  the  heart  and  central  nervous 
system,  the  life  of  the  embryo  depending  upon  it.  The  follow- 
ing illustration  will  serve  for  impressing  the  matter.  The 
arrows  indicate  the  course  of  the  blood  stream.  Only,  the 
diagram  is  misleading  in  several  essential  points,  which  I  am 
compelled  to  notice,  but  it  grieves  me  to  have  it  to  do  ;  nota- 
bly :  1.  The  ductus  arteriosus  (6)  is  nearly  as  large  as  the 
umbilical  vein  itself  (3),  creating  the  impression  that  the 
greater  portion  of  the  umbilical  blood  continues  directly  on 
into  the  vena  cava,  whereas  fully  three-fourths  is  diverted 
into  the  liver  ;  therefore  tending  to  misdirection,  doing  harm 
to  the  medical  pupil,  and  retarding  physiological  progress 
as  well.  2.  The  portal  vein  (7)  is  too  large  in  propor- 
tion— is  larger  than  the  combined  umbilical  and  portal 
streams  (4,  4),  and  is  made  to  appear  as  though  it  debouches  in 
the  umbilical  vein  with  the  right  hepatic  vein,  thence  through 
the  ductus  venosus  into  the  lower  cava ;  whereas,  it  blends 
with  the  umbilical  stream  at  the  portal  gate,  divides  and 
subdivides  in  the  liver  parenchyma  as  an  artery,  the  um- 
bilical and  the  portal  blood  forming  the    common    stream, 


442 


OCCASION  FOR  A   DUCTUS   VENOSUS. 


O     Q 


u  u 


Fig.  186. — Diagram  of  the  Fatal  Circulation.  1,  The  umbilical  cord,  consisting  of  the 
umbilical  vein  and  two  umbilical  arteries,  proceeding  from  the  placenta  (2)  ;  3,  the 
umbilical  vein  dividing  into  three  branches  ;  two  (4,  4)  to  be  distributed  to  the  liver, 
and  one  (5) ,  the  ductus  venosus,  which  enters  the  inferior  vena  cava  (6)  ;  7,  the 
portal  vein,  returning  the  blood  from  the  intestines,  and  uniting  with  the  right 
hepatic  branch  ;  8,  the  right  auricle  ;  the  course  of  the  blood  is  denoted  by  the  arrow 
proceeding  from  8  to  9,  the  left  auricle  ;  10,  the  left  ventricle  ;  the  blood  following 
the  arrow  to  the  arch  of  the  aorta  (11),  to  be  distributed  through  the  branches  given 
oS:  by  the  arch  to  the  head  and  upper  extremities.  The  arrows  12  and  13  represent 
the  return  of  the  blood  from  the  head  and  upper  extremities  through  the  jugular  and 
subclavian  veins,  to  the  superior  vena  cava  (14),  to  the  right  auricle  (8),  and  in  the 
course  oi  the  arrow  through  the  right  ventricle  (15),  to  the  pulmonary  artery  (16)  ; 
17,  the  du  'tus  arteriosus,  which  appears  to  be  a  proper  continuation  of  the  pulmonary 
artery  ;  the  offsets  at  each  side  are  the  right  and  left  pulmonary  arteries  cut  off. 
The  ductus  arteriosus  joins  the  descending  aorta  (18,  18),  which  divides  into  the 
common  iliacs,  and  these  into  the  internal  iliacs,  which  become  the  umbilical  arteries 
(19),  and  return  the  blood  along  the  umbilical  cord  to  the  placenta,  and  the  external 
iliacs  (20),  which  are  continued  into  the  lower  extremities.  The  arrows  at  the  termi- 
nation of  these  vessels  mark  the  return  of  the  venous  blood  by  the  veins  to  the  inferior 


OCCASION   FOR   A   DUCTUS   VEI^OSUS.  443 

while  the  ductus  venosus  conveys  the  unmixed  arterial 
"blood  to  the  lower  cava.  But  the  principal  objection,  that 
the  hepatic  blood  is  not  delivered  into  the  vena  cava  at 
all,  the  hepatic  veins  not  being  represented  by  any  circum- 
stance in  the  diagram.  And  to  call  this  "the  foetal  circula- 
tion," with  fully  three-fourths  of  the  blood  cut  off,  is  simply 
absurd.  It  is  like  knocking  the  bottom  out  of  the  boat  to  give 
it  a  neat  appearance ;  or,  rather,  constructing  a  boat  with- 
out a  "  bottom,"  everything  taut  and  ship-shape  but  the 
bottom,  which  is  absent.  If  physiology  is  to  advance,  it  will 
uever  be  over  that  line.  The  anxiety  to  show  the  wonderful 
double  spiral  twist  in  the  blood  stream  within  the  foetal  heart, 
by  means  of  which  the  blood  from  the  lower  cava  is  transferred 
to  the  ascending  aorta  (8,  9,)  through  the  foramen  ovale,  etc., 
and  that  from  the  upper  to  the  descending  aorta  through  the 
ductus  venosus  (16,  17),  I  conceive  to  be  the  explanation  of 
the  great  blunder.  What  is  needed,  above  all  things,  are  the 
anatomical  facts ,  and  the  special  law  which  applies  in  the 
case  ;  and  by  regarding  anatomy  from  the  stand-point  in  the 
organic  laws,  physiology  is  placed  upon  firm  ground  and  the 
dismal  morass  made  fertile  soil. 


CHAPTER  XVIII. 

DUALISM   IX   MUSCLES   AND   NERVOUS   FORCE. 

Nature  of  Vital  Force — Principle  in  Expansion  and  Contraction — Molecular  Changes  in 
the  Cell-Contents  Involved  in  both  Movements  ;  Illustrated  by  the  Action  in  Pro- 
toplasm and  in  Muscle  Cells  during  Contraction,  as  Seen  in  the  Field  of  the  Polar- 
izing Microscope — Dualism  in  Nervous  Force  Essential  to  the  Production  of  both 
Movements — Extraordinary  Hardness  Produced  in  the  Muscles  by  Nervous  Fore© — 
Explanation  Therefor — Hardness  a  Measure  of  Work — Mode  of  Demonstrating  This 
Circumstance — An  Easy  Method  of  Proving  Dualism  in  Muscles,  and  by  Implication 
Xervous  Force — Lessons  Taught  by  the  Phenomena  in  the  Leech — Ditto,  Tongue  of 
the  Frog — Ditto,  Tortoise  ;  the  Action  in  the  Head,  Neck  and  Tail  Demonstrating 
Dualism  in  Muscles  and  Nervous  Force — Ditto,  Conchifera,  for  Opening  and  Clos- 
ing the  Valves  ;  Together  with  Physiological  Experiment  Demonstrating  the  Cir- 
cumstance— Ditto,  Inferior  Maxilla  in  the  Dog,  showing  the  Masseter  and  Tem- 
poralis Muscles  are  Operated  in  the  Same  Way,  the  Mouth  being  Opened  and 
Closed  by  Means  of  Expansion  and  Contraction  in  these  Muscles — Physiological 
Experiment  upon  the  Nerves  to  the  Ciliary  Ganglion  Proving  Dualism  in  Nervous 
Force  upon  the  Nerves — The  Circumstance  Applied  to  the  Oral  Muscles  and  the 
Action  in  all  the  Sphincters,  all  of  Them  in  Common  Possessing  Circular  and 
Radiating  Muscles,  the  Same  as  the  Iris — Action  in  Erectile  Tissue  Readily  Ex- 
plained ;  Elucidated  by  the  Action  in  the  Tongue  of  the  Chameleon,  Penis,  etc. — 
The  Special  Role  in  Nerves  with  Respect  to  Nervous  Currents,  Nervous  Centres, 
Separators  and  Delimitators  of  Electrical  Fluids  Generated  in  the  Tissues  and 
Carried  to  the  Centres  for  this  Purpose — Reasons  Therefor. 

In  order  to  complete  the  argument,  it  will  now  be  necessary 
to  enter  a  little  more  fully  into  the  mechanics  for  producing 
expansion  and  contraction  in  the  tissues  and  organs,  bringing 
into  prominence  the  principle  in  animal  movement,  making 
this  also  clear  and  easily  understood.  Nor  will  it  be  necessary 
to  unduly  expand  the  argument,  exceeding  the  limits  allotted 
to  this  work,  since  we  are  dealing  with  principles,  and  a  few 
illustrative  cases  will  be  sufficient  for  the  purpose,  at  the  same 
time  maintaining  continuity  throughout,  the  whole  resting 
upon  the  action  in  the  polar  forces  intensified  by  electrical 
currents.  Thus,  we  have  seen  (p.  6)  that  there  is  but  one 
means  known  to  science  for  suspending  matter  in  space  and 
producing  movements  in  it  contrary  to  gravitation,  which  is 


NATURE    OF   VITAL    FORCE.  445 

by  the  action  of  electricity  and  polar  forces  which,  electricity 
serves  to  intensify,  as  illustrated  by  the  rubbed  glass  and 
silver  leaflet  (Fig.  2).  Vital  Force !  Never  mind  about  vital 
force  ;  one  thing  at  a  time  ;  vital  force  has  had  its  day.  What 
we  positively  know  is,  that  heat  and  electricity  produce  move- 
ments in  matter  contrary  to  gravitation,  and  that  polar  force  is 
the  guiding  principle,  serving  also  to  lock  the  molecules  in  the 
positions  which  they  occupy  in  the  structures,  animate  or  in- 
animate, as  the  case  may  be ;  at  the  same  time  maintaining 
mobility  so  as  to  produce  the  requisite  changes  among  the 
molecules  involved  in  the  varied  animal  movements,  having 
its  acme  in  the  muscular  and  nervous  structures,  while  this 
in  turn  has  its  culminating  point  in  warm-blooded  animals  in 
which  body -temperature  is  highest.  Accepting  this,  which  is 
incontrovertible,  all  we  have  to  do,  then,  is  to  make  inspec- 
tion of  the  special  molecular  physics,  since  it  is  by  means 
of  the  changes  effected  in  the  relative  positions  of  the  mole- 
cules in  the  cell-contents  that  the  changes  of  form  are  pro- 
duced in  the  cells  and  in  the  mass  composing  the  muscles, 
etc.,  whereby  the  movements,  voluntary  and  involuntary, 
are  produced,  the  whole  resting  upon  molecular  physics  and 
the  action  of  the  polar  forces,  which  heat  and  electricity  serve 
to  intensify.  And  that  this  is  also  true,  is  fully  proven  by  the 
circumstance  that  by  simply  reducing  temperature,  the  move- 
ments are  promptly  arrested,  while  heat  undergoes  metamor- 
phosis into  electrical  force  for  energizing  the  polar  forces,  the 
same  as  in  air  and  water,  producing  the  vital  phenomena. 
There  !  that  is  your  Vital  Force. 

Commencing  at  the  bottom,  so  to  speak,  of  this  problem,  we 
have,  then,  the  action  in  protoplasm  to  begin  with.  In  undif- 
ferentiated protoplasm  no  form- elements  present ;  nevertheless, 
this  animal  (Moneron)  is  highly  sensitive,  responds  quickly  to 
irritants,  performs  voluntary  movements,  eats,  respires,  circu- 
lates its  fluids,  and  reproduces  itself,  the  form-elements  (flrst 
nucleus)  in  the  subsequent  stages,  or  amoeboid  development, 
the  muscles  and  nerves  in  the  compound  organisms  for  ener- 
gizing the  local  actions  and  effecting  coordination,  that  the 
whole  may  perform  as  but  a  single  individual  only,  the  vascu- 
lar system,  respiratory  apparatus  and  the  other  organs  undergo- 


440  NATTRE    OF   VITAL    FORCE. 

ing  commensurate  changes  with,  the  stage  in  development,  there 
being  a  progressive  onward  movement  in  this  direction  ;  while 
the  whole  relates  to  perfection  of  mechanical  work  for  evolving 
the  force  which  is  expended  in  them  in  producing  the  varied 
movements,  while  this  in  turn  relates  to  the  struggle  with 
gravitation,  which  is  pivotal  in  development ;  but  coexistent 
with  this,  the  law  of  pressure,  which  is  being  incessantly 
invoked  by  means  of  the  rhythmical  expansions  and  contrac- 
tions taking  place  in  the  organs  and  tissues  for  compelling 
circulation  in  the  measure  of  the  physiological  requirements, 
which  is  also  produced  by  means  of  heat  and  electricity,  the 
same  as  for  producing  the  voluntary  movements  and  the 
actions  in  air  and  water.  And  if  you  would  find  Vital  •Force, 
you  would  have  to  seek  for  it  in  heat-metamorphosis,  with  the 
sun  as  the  common  source  of  supply.  The  "  Fire- Worshiper" 
had  instinctively  sought  out  this  circumstance  long  since.  And 
"he  laughs  best  who  laughs  last."  Notwithstanding,  the  evi- 
dence of  an  all-pervading  Intelligence  is  incontrovertible. 

For  impressing  the  circumstance  of  molecular  action,  we 
first  bring  before  the  reader  the  following  forcible  illustration 
(Fig.  187),  by  the  famous  biologist  at  Jena,  representing  eight 
presentations  of  the  same  animal  (blood-cell  of  a  snail),  at  as 
many  moments  during  feeding.  For  example,  beginning  with 
./,  we  have  the  spherical  and  more  or  less  translucent  condition 
the  cell  at  first  presents,  then  an  extension  of  the  branched 
processes  (b)  for  pumping  the  fluids  into  the  body,  else  flow- 
ing over  them  and  so  engulfing  the  food  particles,  then  the 
conditions  at  c,  d,  e,  f,  g,  h,  all  the  while  becoming  more 
and  more  opaque  from  absorption  of  the  food,  till  at  last  the 
condition  is  reached  when  no  more  food  can  be  taken,-  the  ani- 
mal being  full,  as  in  the  case  of  the  sucking  leech  (Figs.  1 5-17), 
and  for  every  stage  in  development,  the  corresponding  increase 
in  size  being  alike  visible  in  all ;  notably  h  is  much  larger  than 
a,  the  full  leech  (Fig.  17)  than  the  empty  (Fig.  15).  Now, 
then,  in  order  to  produce  these  varied  changes  of  form  in  the 
cell,  it  is  manifest  that  the  cell-contents,  the  molecules  equally 
with  the  masses,  would  have  to  change  their  relative  positions, 
even  to  the  nucleus  (a,  o),  which  occupies  the  root  of  a  large 
branched  process  in  g  ;   while  in  the  subsequent  condition  (h), 


MOLECULAR   MOVEMENTS    IN   CELLS. 


447 


in  which  the  animal  is  fully  contracted,  restoring  the  original 
spheroidal  condition  (a),  the  nucleus  occupies  a  nearly  central 
position.  In  other  words,  we  have  definite  actions  and  limita- 
tions in  the  protoplasmic  substance,  the  movements  directed 
to  special  objects,  whether  it  relate  to  feeding  or  locomotion, 
in  the  entire  absence  of  muscles  or  nerves,  or  any  form-elements 
for  producing  them.  Now,  then,  by  looking  from  this  to  the 
actions  taking  place  in  the  cells  of  compound  organisms,  in 
which  muscles  and  nerves  are  developed,  it  will  at  once  be 
seen  that  the  same  principle  for  producing  motion  is  main- 
tained, and  must  be  so  in  the  very  nature  of  things.     In  the 


Fig.  187. — Devouring  Blood-Cells  of  a  Naked  Sea-Snail  (Thetis)  very  much  magnified. 
In  connection  with  the  blood-cells  of  this  snail,  I  was  the  first  to  observe  the  impor- 
tant fact  that  "  the  blood-cells  of  invertebrate  animals  are  uncovered  lumps  of  proto- 
plasm, and,  like  the  Amoebae,  by  means  of  their  peculiar  movements  can  absorb 
matter,"  can,  therefore,  "eat."  When  at  Naples  (on  the  10th  of  May,  1859)  I  had 
injected  the  blood-vessels  of  one  of  these  snails  with  pulverized  indigo  dissolved  in 
water.  I  was  much  astonished  to  find  after  a  few  hours' that  the  blood-cells  them- 
selves were  more  or  less  filled  with  fine  particles  of  indigo.  By  repeated  experi- 
mental injections,  I  was  able  to  watch  the  absorption  of  the  coloring  matter  into  the 
blood-cells,  which  was  accomplished  exactly  as  by  Amoebse.  (See  "Monograph  of 
Radiolaria,"  1862,  pp.  104,  105.)— Haeckel. 

muscles,  for  example,  in  which  changes  of  form  in  the  muscle 
cells,  produced  by  the  molecular  movements  in  the  cell- walls 
and  cell -contents,  are  referable  the  expansile  and  contractile 
actions  in  the  muscles,  in  response  to  nervous  force ;  while  the 
very  fact  that  nervous  force  exerts  this  effect  upon  the  mole- 
cules, shows  at  once  it  is  also  electrical,  thereby  producing 
the  changes  in  polarity  in  the  molecules,  which  is  foundation 
and  source  of  all  the  movements  in  animate  as  well  as  in  in- 
animate nature.  In  fine,  work  is  produced  in  muscles  and 
animal  tissues  by  means  of  heat  metamorphosis  and  the  action 
of  electrical  force,  or  the  same  as  obtains  in  any  mechanism 
in  motion,  only  that  in  animal  life  it  is  automatic  ;  neverthe- 
less, every  movement  must  have  its  equivalent  in  force,  be  paid 


448  i  NATURE   OF  VITAL   FORCE. 

for  at  the  time,  there  and  then,  else  there  could  be  no  move- 
ment, consequently  no  life. 

Furthermore,  we  can  readily  understand  why  the  pumping 
actions  for  increasing  circulation  should  correspond  with  the 
activities,  rising  and  falling  with  these,  as  everything  is  paid 
for  by  circulation,  and  this  could  not  be  made  commensurate  in 
any  other  way,  pressure  being  invoked  by  the  organs  in  the 
measure  of  the  requirements  for  compelling  prompt  response, 
while  for  producing  this  it  also  calls  for  heat  and  electrical 
force,  so  that  the  principle  is  at  once  seen  to  be  all-pervading. 
By  reducing  temperature  simply,  the  action  is  promptly 
arrested  in  both  animal  and  floral  life.  Thus,  water  (which 
is  chief  constituent  of  the  tissues,  and  may  be  roughly 
estimated  at  three-fourths,  the  gray  substance  of  the  brain 
80.5  decimals  in  the  100)  freezes  at  a  temperature  of  30  de- 
grees Fahr.,  which  would  at  once  put  an  end  to  circulation, 
therefore  of  life  ;  but  long  before  this  reduction  in  tempera- 
ture is  produced  in  the  tissues,  all  visible  phenomena  have 
arrest,  which  establishes  beyond  question  that  heat  is  the 
agent  for  effecting  the  actions.  Not  that  heat  is  actually  life 
itself,  but  that  life  is  product  of  heat  metamorphosis,  and  the 
interaction  of  the  polar  forces  in  special  forms  of  matter,  as 
embraced  in  air  and  water,  out  of  which  the  living  organism 
is  evolved. 

Finally,  continuity  (which  undoubtedly  exists)  implies  simi- 
larity in  force,  so  that  in  every  aspect  of  the  case  we  see  neces- 
sity for  proceeding  upon  this  basis,  in  order  to  make  the 
matter  intelligible.  And  since  force  connects  through  and 
through  in  order  to  produce  the  actions  for  sustaining  life, 
nowhere  more  visible  than  in  animal  life,  for  producing  the 
movements  in  respiration,  circulation  and  the  voluntary  ac- 
tions, it  follows  that  there  should  be  some  common  principle, 
applicable  alike  to  all,  for  effecting  *the  molecular  actions  in- 
volved in  them,  and  which  we  find  to  be  the  polar  forces,  or 
the  same  as  for  producing  the  actions  in  air  and  water ; 
otherwise  is  inexplicable.  In  furtherance  of  this  view,  we 
have  submitted  the  action  in  protoplasm,  showing  it  is  molec- 
ular ;  but  we  would  now  contrast  this  with  the  action  taking 
place  in  muscle  cells  during  contraction,  showing  it  is  equally 


expansion  and  contraction.  449 

applicable  to  this  also,  as  the  following  forcible  excerpt,  by  a 
distinguished  German  microscopist,  clearly  establishes  :* 

"An  important  question  still  remains,  which  can  be  solved 
by  the  help  of  the  polarizing  microscope :  Are  the  sarcous  ele- 
ments to  be  regarded  as  single  and  individual  elementary 
bodies,  or  as  groups  of  solid  bodies,  capable  of  being  variously 
disposed  ?  If  the  muscles  contract,  the  fibres  are  seen  to  be- 
come thicker,  and  the  transverse  striae  to  approximate.  Each 
sarcous  element  must  consequently  change  its  form  and  be- 
come shorter  and  thicker.  If  such  a  change  of  form  result 
from  any  force  acting  in  an  elementary  solid  body,  the  opera- 
tion of  the  force  must  extend  as  far  as  the  individual  mole- 
cules, the  optic  constants  must  be  changed,  and  it  is  nob  con- 
ceivable that  they  should  be  so  changed  that  the  ordinary  and 
the  extraordinary  ray,  after  they  have  traversed  equal  thick- 
nesses in  the  same  direction,  should  present  again  the  same 
difference  in  velocity  that  they  offered  under  similar  circum- 
stances before  the  change  of  form.  But  it  is  quite  a  different 
matter  if  the  sarcous  elements  are  groups  of  solid,  doubly  re- 
fracting bodies,  of  which  each  individual  remains  unchanged 
in  form  in  the  act  of  contraction.  The  form  of  the  whole 
group — that  is,  of  the  sarcous  element — is  here  changed  by  an 
alteration  in  the  arrangement  of  the  several  corpuscles,  just  as 
in  a  company  of  soldiers,  groups  of  various  breadths  and 
depths  are  produced  by  changes  in  the  position  of  the»several 
individuals.  In  the  latter  case,  the  optic  constants  are  not 
altered  in  the  act  of  contraction,  and  the  rays,  on  this  account, 
if  they  have  traversed  equal  thicknesses  in  the  same  direction, 
must  constantly  exhibit  the  same  differences  in  velocity, 
whether  the  muscles  be  in  the  relaxed  or  in  the  contracted 
condition.  Since  we  have  a  measure  of  the  difference  of  veloc- 
ity in  the  colors  which  appear  under  the  polarizing  micro- 
scope, we  are  enabled  to  answer  the  question  experimentally 
whether  the  optic  constants  of  the  contractile  substance  change 
during  contraction  to  any  considerable  extent  or  not.  All  the 
investigations  I  have  directed  to  this  point  have  had  a  nega- 
tive result — i.  e.,  I  have  never  seen  any  alteration  of  color  that 

*  The  Behavior  of  Muscular  Fibres  when  Examined  by  Polarized  Light. 
By  E.  Bnicke      Strieker's  Manual  of  Histology. 


460  EXPANSION   AND   CONTKACTION. 

could  not  be  entirely  referred  either  to  changes  in  the  thick- 
ness of  the  layer  traversed,  or  in  the  angle  which  the  rays  un- 
dergoing interference  make  with  the  optic  axis.  As,  there- 
fore, I  have  in  vain  sought  after  a  change  of  the  optic 
constants,  I  must  maintain  that  the  sarcous  elements  are  not 
elementary  and  simple  solid  bodies,  but  groups  of  smaller, 
doubly  retractile  bodies." 

This  brings  the  molecular  physics  within  the  range  of 
thought  and  distinct  mental  presentation.  But  then,  the  dis- 
tinguished author,  it  would  seem  to  us,  has  omitted  one-half 
of  the  mechanics,  namely,  expansion  ;  for  expansion  and  con- 
traction are  correlated  forces.  And  keeping  expansion  for 
the  time  in  view  only,  is  it  not  as  easy  to  perceive  that  it  may 
be  made  the  primary  movement,  the  individual  molecules 
going  from  the  middle  to  the  ends  of  the  column,  in  order  to 
effect  it,  then,  by  simply  reversing  the  movement,  produce 
contraction  \ 

In  point  of  fact,  it  necessitates  as  much  discipline  among  the 
molecules  in  reversing  the  action  as  for  producing  the  primary 
movement ;  this,  whether  it  relates  to  contraction  or  expansion. 
In  short,  that  the  molecules  may  go  through  either  evolu- 
tion with  equal  ease  and  celerity,  in  obedience  to  special  ner- 
vous stimulus  which  applies  for  effecting  it.  There  would 
seem  to  be  no  reason  why  this  may  not  be  the  case.  Given 
that  the  molecules  are  held  in  their  relative  positions  by  the 
interaction  of  the  polar  forces  (of  which  there  is  no  room  for 
doubt,  since  it  applies  to  everything),  it  follows  that  dualism 
inheres  in  the  mechanics,  while  movement  is  product  of  a 
change  in  polarity  among  the  molecules.  Furthermore,  this 
principle  in  the  mechanics  would  afford  a  ready  explanation 
for  a  number  of  phenomena  otherwise  inexplicable,  notably : 

a.  How  simultaneous  action  may  be  produced  in  opposite 
sets  of  muscles — e.  g. ,  flexors  and  extensors  with  the  obviation 
of  friction  and  a  waste  of  force,  to  the  end  that  each  should 
perform  an  active  part  in  the  work  to  be  accomplished,  and 
not  hindering  but  aiding  each  other.  But  at  present  one  set 
is  made  to  pull  the  other  into  extension,  which  is  unscientific, 
since  it  involves  destruction  or  loss  of  equilibrium  upon  which 
the  mechanics  is  based,   not  to  mention  rude  friction  and 


EXPANSION   AND   CONTRACTION.  451 

waste  of  force  thereby  made  inevitable.  In  place  of  this,  how- 
ever, we  have  both  sets  of  muscles  acting  simultaneonsly  and 
in  perfect  concert  with  each  other,  while  equilibrium  is  main- 
tained in  this  manner  at  any  angle  in  flexion  or  extension,  at 
the  same  time  obviating  strain  and  friction  to  the  histological 
elements  as  well  as  the  articular  cartilages  in  the  joints,  other- 
wise inevitable.  In  regard  to  the  latter,  for  example,,  we  may 
take  the  action  in  the  hand.  Spread  it  open  widely,  then 
tightly  close  it  as  a  fist.  How  swiftly  this  is  done!  Well, 
is  the  enormous  extension  of  the  flexors  in  the  first  due  to 
contraction  in  the  opposing  extensor  muscles?  and  in  the 
second  is  the  prodigious  extension  of  the  extensors  due  to 
contraction  in  the  flexors  ?  In  this  mechanics  we  would  have 
the  ends  of  the  fingers  in  which  the  tendons  are  inserted  -as 
the  point  oVappui  from  which  force  would  have  to  be  exerted 
in  either  case,  and  during  closing  of  the  hand  the  articular 
cartilages  would  have  to  endure  rude  friction  from  the  rub- 
bing of  the  opposing  tendons  over  them,  increasing  pari  passu 
with  the  movement,  so  that  when  the  hand  is  tightly  closed 
the  pressure  upon  the  articular  ends  of  the  bones  would 
necessarily  be  enormous,  painful  even  to  contemplate  ;  the  same 
remark  applying,  of  course,  for  extension,  as  the  strain  would 
have  to  fall  upon  the  articular  cartilages  in  either  case.  On  the 
contrary,  however,  we  are  not  conscious  of  any  strain  or  fric- 
tion whatever,  either  in  the  ends  of  the  digits  or  the  articular 
ends  of  the  bones,  which  shows  conclusively  that  one  set 
of  muscles  is  not  pulled  into  extension  by  the  other,  but  that 
both  act  together  and  simultaneously,  the  one  expanding  as 
the  other  is  contracting,  and  vice  versa. 

b.  But  we  have  now  to  mention  another  important  factor  con- 
nected with  musculation,  which  is  also  needing  explanation, 
notably  the  extraordinary  hardness  and  firmness  produced  in 
the  muscles  by  nervous  force  during  musculation,  and  increas- 
ing with  the  energy  in  expansion  and  contraction,  since  it  ap- 
plies equally  to  both  sets  of  muscles,  to  flexors  as  well  as 
extensors,  which  are  simultaneously  affected.  The  following 
illustration  (Fig.  188)  will  serve  for  impressing  the  matter.  And 
with  the  weight  in  the.  body  thus  resting  upon  the  metatarsal 
bones,  the  foot  fully  extended,  both  sets  of  muscles  to  palpation 


452  ELECTKICAL    TENSION". 

seem  as  hard  as  iron,  so  great  is  the  hardness.  Furthermore, 
the  hardness  is  in  the  ratio,  of  the  weight  sustained.  For 
example,  if  the  foot  is  extended  when  sitting  or  lying  down, 
the  muscles  show  only  an  amount  of  hardness ;  but  if  a 
portion  of  the  body-weight  be  then  placed  upon  it,  at  once 
there  is  greater  hardness,  while  this  increases  in  the  ratio 
of  the  weight,  till  finally  the  maximum  is  reached,  when 


!Fig.  188. — Showing  the  great  hardness  which  is  produced  in  the  muscles  by  nervous 
force  when  in  energetic  action,  and  which  applies  to  both  flexor  and  extensor 
muscles,  proving  the  dualism  in  nervous  force,  or  the  power  of  producing  siniul- 
taucous  contraction  and  expansion  in  opposite  sets  of  muscles,  coordination  being 
perfect. 

the  whole  weight  in  the  body  is  thrown  upon  it ;  pro- 
vided, of  course,  the  party  can  endure  no  more  than  this  ; 
otherwise  it  will  continue  to  swell  till  the  end  of  his  physical 
powers  is  reached,  for  the  hardness  will  correspond  with  the 
amount  of  work  the  muscles  are  performing. 

Taking  work,  then,  as  the  condition  for  producing  hardness, 
it  follows  that  both  sets  of  muscles,  flexors  with  extensors  per- 
form work  during  musculation.  The  great  advantage  of  this 
is,  that  while  it  maintains  equilibrium,  thereby  reducing  fric- 


ELECTKICAL    TENSION.  453 

tion  and  conserving  force  and  structure,  it  imparts  great  firm- 
ness to  the  articulations,  the  tendons  acting  as  "splints"  upon 
the  joints,  since  the  hardness  extends  to  them  as  well.  It  is, 
therefore,  obvious  that  one  may  not  speak  of  "relaxation"  at 
all  in  connection  with  musculation,  being  as  wide  apart  as  the 
' '  poles' '  — antipodes. 

In  short,  "relaxation"  is  a  state  of  rest,  the  very  opposite 
to  work  ;  at  the  same  time,  it  also  involves  dualism,  which  is 
essential  to  equilibrium,  for  which  purpose  the  limbs  are 
placed  at  an  angle  between  flexion  and  extension,  therefore 
involving  action  in  both  sets  of  muscles  in  order  to  effect 
it ;  hence,  the.  monistic  theory  of  muscular  force  will  not 
apply  here  either.  Moreover,  the  crucial  test  of  a  theory 
is  its  power  of  absorbing  all  the  phenomena.  It  comes 
to  this,  that  the  nervous  apparatus  itself  is  based  upon  the 
molecular  or  polar  actions,  its  special  office  being  to  energize 
the  polar  forces  for  increasing  the  action  in  the  organs  com- 
mensurate with  the  exigencies  in  the  functions,  and  for  effect- 
ing coordination,  to  the  end  that  a  balance  may  be  maintained 
in  the  organism,  the  whole  performing  as  but  a  single  indi- 
vidual. 

In  Reference  to  the  Cause  of  the  Hardness  in  the  Muscles, 
which  is  Produced  by  Nervous  Force,  and  Increasing  in 
the  Measure  of  This. 

In  casting  about  for  a  natural  solution  of  the  deeply  inter- 
esting problem  in  musculation — notably,  the  great  hardness 
and  firmness  produced  in  the  muscles  by  nervous  force — we  are 
struck  with  the  remarkable  experience  of  Faraday  with  the 
electro-magnet,  which,  for  several  reasons,  would  seem  to  us  to 
afford  a  natural  and  easy  solution  of  the  phenomenon  ;  more- 
over, it  is  in  entire  accordance  with  the  fundamental  principle 
in  the  mechanics — i.  e.,  the  interaction  of  the  polar  forces, 
intensified  by  electrical  currents.  For  example,  he  discovered 
when  the  current  was  on  in  an  electro-magnetic  machine,  that 
"a  weight  of  several  pounds  of  copper,  when  made  to  move 
through  the  magnetic  field,  though  nothing  is  visible,  appears 
to  move  through  a  viscous  fluid  ;  while,  when  a  flat  piece  of  the 
metal  is  caused  to  pass  to  and  fro  like  a  saw  between  the  poles, 
the  sawing  of  the  magnetic  field  resembles  the  cutting  through 


454  MOLECULAK   ACTION    AND     ELECTEICTTY. 

of  cheese  or  butter."  And  by  connecting  this  deeply  suggestive 
fact  with  that  more  recently  ascertained  in  physiology  (Helm- 
holtz,  Holmgren,  Burdon-Sanderson),  namely,  that  electrical 
change  precedes  mechanical  change  in  the  sarcous  elements, 
the  hardness  in  the  muscles  which  accompanies  musculation 
increasing  with  the  afflux  of 'nervous  force  in  the  parts,  would 
seem  to  us  to  be  due  to  a  similar  cause,  or  the  interaction  of 
the  polar  forces  intensified  by  electrical  currents.  In  other 
words,  that  the  hardness  represents  magnetic  tension,  pro- 
duced by  electrical  force,  and  that  the  nervous  currents  but  a 
form  of  electricity,  of  which  there  can  be  but  little  doubt,  con- 
tinuity in  force  for  producing  molecular  action  calling  for  this. 
Furthermore,  it  would  afford  an  explanation  of  other  phenomena, 
otherwise  inexplicable  ;  notably  :  a.  How  the  worms  can  erect 
their  soft,  tubular  bodies  almost  perpendicularly  with  the 
utmost  ease  and  celerity,  when  climbing  and  locomoting.  b.  It 
would  afford  an  explanation  for  the  extraordinary  feats  of 
strength  displayed  during  periods  of  excitement  sufficient  to 
tear  the  muscles  apart  when  out  of  the  body,  since  the  increase 
in  magnetic  tension  in  the  sarcous  elements  should  increase 
cohesive  power  correspondingly  ;  in  this  way  subserving  the 
highest  uses  in  the  organism  by  reducing  strain,  thereby  con- 
serving structure  and  increasing  function  at  one  and  the  same 
time.  For  the  muscle  elements,  being  thus  sustained  by  elec- 
trical force,  and  operated  by  electrical  force,  friction  and  strain 
during  musculation  are  made  impossible,  a  perpetual  balance 
in  force,  which  it  necessarily  involves,  making  it  impossible. 
And,  what  is  of  equal  importance,  action  is  thereby  unified,  all 
the  parts  moving  together  in  perfect  order  under  electrical 
force,  which  pervades  the  mechanics  through  and  through, 
producing  the  utmost  concert  in  action. 

Having  thus  gone  briefly  over  the  mechanical  principles  in 
musculation,  we  are  now  prepared  to  take  up  special  vital 
phenomena  appertaining  to  musculation  and  circulation,  hither- 
to inexplicable,  the  first  in  order  being  the  demonstration  of 
dualism  in  the  muscles,  which  is  readily  done. 

An  Easy  Mode  of  Demonstrating  Dualism  in  the  Muscles. 

For  this  purpose  we  make  selection  from  the  worms,  since 

there  is  no  occasion  for  isolating  the  muscular  fibres  in  them, 


DUALISM     DEMONSTRATED.  455 

the  special  arrangements  which  obtain  making  this  unneces- 
sary, for  the  muscles  being  disposed  in  circular  and  longitudi- 
nal layers,  the  changes  of  form  which  the  body  undergoes  is 
sufficient  in  itself  to  indicate  the  action  in  the  muscles,  while 
it  gives  us  the  actual  facts  in  musculation,  which  the  isolated 
fibre,  removed  from  the  cycle  of  organic  forces  and  the  volition 
of  the  animal,  cannot  possibly  do. 

Furthermore,  to  reproduce  the  phenomena  in  musculation 
requires  perfect  knowledge  of  the  special  mechanics  and  the 
power  on  the  part  of  the  operator  to  isolate  the  positive  and 
negative  electricities  ;  at  the  same  time,  to  so  regulate  the  cur- 
rents as  not  to  exceed  the  normal  limits,  which,  it  is  needless 
to  add,  is  difficult  of  attainment  ;  nevertheless,  the  power  on 
the  part  of  the  animal  to  do  this  may  not  be  doubted  for  a 
single  moment,  since  the  matter  is  fully  proven  by  its  ability 
to  produce  simultaneous  expansion  in  opposite  sets  of  muscles, 
and  even  in  different  portions  of  the  individual  muscle  and 
muscular  fibre,  of  which  we  shall  give  ample  proof  before  we 
are  done,  commencing  with  the  worms,  notably  the  leech  (Fig. 
189),  and  ending  with  the  higher  animals.  Here,  of  course,  we 
must  begin  with  the  anatomy  as  means  to  ends,  being  the 
arrangements  for  producing  worlc,  while  the  special  actions  or 
movements  observable,  together  with  the  laws  of  pressure  and 
gravitation  which  apply,  and  the  dual  principle  in  nervous 
force  for  producing  the  molecular  movements,  will  give  us  the 
method  of  doing  this,  which  is  what  we  wish  to  get  at  in 
the  case.  In  short,  whether  there  is  actually  dualism  in 
muscles. 

Anatomy. — Fundamentally,  the  body  of  the  worms  is  a  hol- 
low cylinder,  composed  of  circular  and  longitudinal  muscular 
layers,  the  former  external,  the  latter  internal.  In  Annelides, 
the  longitudinal  fibres  are  increased,  forming  two  dorsal  and 
two  ventral  layers,  leaving  a  lateral  groove.  In  addition  to 
this,  a  layer  of  transverse  fibres,  generally  in  the  form  of  dis- 
tinct bundles,  passes  from  the  ventral  median  line  to  the  lateral 
grooves,  which  enables  the  animal  to  flatten  its  body  when 
swimming,  passing  through  the  water  by  graceful  undulations 
of  the  body,  which  is  flattened  out,  ribbon-like.  The  suckers 
are  special  differentiations  of  the  dermo-muscular  tube,  which 


466 


DUALISM      DKMONSTRATKD. 


agree  with  one  another  in  all  the  essential  points  of  their 
structure. 

The  great  relative  thickness  of  the  muscular  cylinder  is  seen 
in  the  following  illustration  (Fig.  13,  A,  B,  in). 

During  feeding  and  locomotion  the  muscular  layers  are  af- 
fected differently,  the  one  directly  the  opposite  of  the  other, 

B  B  J) 


Fig.  189. — Six  Cuts  of  the  Leech,  showing  as  many  changes  of  form,  assumed  within  a 
few  minutes,  allowing  for  the  time  occupied  in  filling  itself  with  blood,  and  neces- 
sarily involving  simultaneous  expansion  and  contraction  in  the  circular  and  longi- 
tudinal muscles.  A,  before  feeding  ;  B,  when  extending  itself  ;  C,  when  feeding  ; 
D,  when  gorged  with  blood  ;  JE7,  when  freeing  his  "feet"  for  locomotion,  driving 
the  blood  to  the  central  portions  ;  F,  when  locomoting. 

but  acting  harmoniously  and  in  utmost  concert,  their  joint 
action  being  essential  for  effecting  the  movements.  Kow, 
then,  and  taking  the  state  of  rest  as  more  nearly  representing 
the  condition  of  equilibration  where  motion  is  reduced  to  a 
minimum,  we  have  presented  to  us  the  following  appearance 
(Fig.  189,  A),  in  which  the  animal  is  very  small,  nearly  round 


DUALISM     DEMONSTRATED.  457 

or  ovoidal  in  form.  But  when  locomotion  sets  in,  all  this 
is  changed  (B),  and  from  a  short,  globular  body  he  assumes  a 
long  rod-like  shape,  extending  himself  to  more  than  double 
the  length  he  had  when  resting,  while  the  anterior  portions 
are  drawn  out  to  the  dimensions  of  a  bodkin  or  even  less,  so 
very  attenuated  they  become. 

!Now,  then,  in  order  to  effect  this  action,  the  circular  muscles 
must  contract  and  the  longitudinal  elongate  simultaneously; 
otherwise  it  were  utterly  impossible  to  effect  extension,  since 
the  one  is  greatly  reduced  in  length,  the  other  proportion- 
ately increased  ;  hence,  the  former  must  contract  and  the 
latter  expand.  No?  Please  explain,  then,  and  make  us 
understand  how  it  is  done. 

The  short  muscles  contract  and  pull  out  the  long  muscles  ! 
Nothing  of  the  kind.  Nay,  impossible  and  preposterous. 
Look  at  the  anatomy,  and  put  a  little  thought  into  the  matter. 
In  the  first  place,  the  long  muscles  in  the  leech  are  more 
numerous,  therefore  more  powerful  than  the  short  ;  the 
weaker  force,  then,  would  have  to  overcome  the  stronger, 
which  is  not  reasonable.  1.  It  would  involve  prodigious 
strain  to  the  sarcous  elements  in  the  long  muscles  by  being 
thus  forcibly  pulled  to  such  extreme  limits  ;  it  could  not 
be  otherwise,  in  the  very  nature  of  things.  Moreover,  muscles 
will  not  admit  of  much  forcible  extension,  resisting  it 
promptly,  and  the  greater  the  force  applied  the  more  they 
contract,  tearing  in  twain  when  the  limit  is  reached.  2.  It 
would  involve  enormous  waste  of  force,  which  is  antagonistic 
to  conservation  of  energy  and  of  structure,  which  is  funda- 
mental in  the  organism.  Hence  this  rude  mechanics,  born  of 
the  monistic  theory  of  muscular  function,  cannot  possibly  be 
correct.  Fortunately,  however,  an  easy  method  obtains  for  dis- 
proving it  by  actual  demonstration,  notably,  hy feeding  the  ani- 
mal, when  it  becomes  five  or  six  times  the  size  it  had  in  the  empty 
condition  (A,  D).  Now,  then,  by  looking  from  A  to  D,  it  is 
at  once  perceived  that  important  changes  in  the  muscles  have 
taken  place,  notably  that  D  is  much  larger  than  A,  it  is  much 
longer  than  A — indeed,  nearly  twice  its  length,  while  in  circum- 
ference it  is  a  number  of  times  larger.  Hence  both  sets  of 
muscles,  the  transverse  as  well  as  the  longitudinal,  are  elon- 


453  DUALISM     DEMONSTRATED. 

gated.  Which  set,  then,  pulled  out  the  other  set?  The 
"pulling-out  business"  will  not  answer  at  all ;  one  might  as 
well  try  to  lift  himself  by  his  boot-straps.  Last,  but  not  least, 
beneath  muscular  movement  itself  lies  the  law  of  fluid  equilib- 
rium, which  compels  expansion  in  order  to  produce  low 
pressure  within  the  animal ;  otherwise  the  fluids  would  not 
flow  into  the  internal  parts 

So  then,  we  have  both  the  law  for  compelling  movement  in 
the  fluids  and  the  fact  in  this  enlargement  itself,  as  indis- 
putable proofs  of  muscular  expansion.  And  how  can  it 
matter  whether  we  are  able  to  reproduce  expansion  in 
muscles  artificially  or  not  %  The  bald  fact  remains  that  the 
animal,  of  its  own  volition,  does  it  readily  enough,  and  must 
do  so  in  the  very  nature  of  things  ;  that  ends  the  matter  there : 
ttiis,  however,  that  when  the  operator  shall  have  produced 
expansion  as  readily  as  contraction  in  the  muscles  with  the 
method  of  coordinating  the  movements,  the  mystery  in  vital 
force  itself  will  have  been  solved,  and  the  cause  of  these  in- 
numerable "eddies  "  in  the  boundless  ocean  of  Force  laid  bare 
to  the  investigator,  for  Life  will  then  have  been  seen  through 
and  through.  But  since  it  involves  metamorphosis  in  force, 
his  investigations  will  be  brought  to  a  sudden  stand-still  on  the 
threshold  of  that  awesome  fact — the  heart  of  Nature's  God, 
the  unfathomable  and  unnamable  All. 

The  animal  not  only  produces  expansion  as  readily  as  con- 
traction, but  the  two  actions  are  coordinated  in  producing  cir- 
culation and  the  voluntary  movements,  since  the  method  for 
increasing  circulation  by  rhythmical  changes  in  pressure  in- 
volves alternate  expansions  and  contractions  in  the  muscles 
of  the  heart,  vessels,  and  hollow  viscera  for  compelling  move- 
ment in  the  contents,  the  whole  having  adjustment  with 
pressure  and  which  applies  to  the  mouth  as  well  as  any  other 
portion.  Thus,  in  the  case  of  the  sucking  leech  (0),  the  oral 
sucker  is  first  expanded  over  the  cutaneous  surface  {F),  then 
gradually  contracted  for  pulling  up  the  artificial  nipple,  the 
pharynx  at  the  same  time  expanding  for  drawing  it  into  the 
chamber  (c),  where  it  is  incised  by  the  to-and-fro  motion  of  the 
three  serrated  teeth  in  the  pharynx  by  the  alternate  expansions 
and  contractions  in  the  muscles  connecting  with  the  cartil- 


DUALISM     DEMONSTRATED.  459 

ages,  the  enlargement  forming  the  cnpping-apparatns  (c),  at 
the  same  time  working  energetically  for  aspirating  and  pump- 
ing the  fluids  into  the  canal ;  thence  propelled  along  the  tube 
by  alternating  expansions  and  contractions  contiguous  to  each 
other,  as  in  deglutition,  of  which  it  is  archetypal.*  Finally, 
when  gorged  with  blood,  so  that  no  more  can  be  introduced 
(Z>),  the  suction-action  must  cease,  for  the  limit  in  expansion 
is  reached  ;  at  the  same  time,  however,  this  necessitates  the 
power  to  maintain  expansion  for  producing  equilibrium  in 
pressure,  otherwise  contraction,  by  increasing  pressure,  would 
compel  reflux  through  the  oral  orifice,  else  the  blood  would 
escape  by  the  anus,  as  when  the  animal  is  compressed  between 
the  fingers  for  the  purpose  of  "stripping"  him,  which  is  by 
reflux  action  through  the  oral  orifice,  the  terminal  end  of  the 
gut  being  very  small.  Hence,  all  these  movements  involve 
dualism  in  the  muscles,  the  notable  circumstances  being  the 
energy  in  expansion,  and  the  power  of 'maintaining  a  balance, 
whether  it  relate  to  expansion  or  contraction,  and  which  ap- 
plies for  the  voluntary  movements,  as  well  as  for  circulation. 
For  forcible  evidence  of  this  circumstance,  and  the  facility 
with  which  the  action  is  reversed,  compare  D  with  E,  in  which 
the  animal  is  getting  ready  for  locomoting,  by  first  compelling 
the  blood  from  the  terminal  ends  toward  the  central  portions 
of  the  body,  or  behind  the  more  central  diaphragms,  which 
are  expanded  for  detaining  it,  so  as  to  allow  the  feet  to  be  used 
as  in  F.  Here  contraction  is  made  as  energetic  as  expansion 
had  previously  been,  which  is  again  as  suddenly  reversed 
lor  expanding  the  feet,  which  are  widely  expanded  to  function 
as  the  basis  of  support  in  locomotion ;  while  to  this  again 
must  be  added  the  expansion  which  takes  place  in  the  posterior 
portions  of  the  body,  contiguous  to  the  hind  foot,  for  bringing 
the  blood  over  the  centre  of  gravity  (F),  thereby  relieving  the 
anterior  portions  correspondingly  for  assisting  locomotion, 
enabling  the  animal  to  more  readily  elevate  these  portions  in 
locomotion,  the  ends  being  raised  and  lowered  alternately. 
But  by  reason  of  coagulation  of  the  blood,  which  soon  sets  in, 
and  the  diversion  of  nervous  force  to  the  alimentary  canal 

*  In  this  case,  gravitation  aids  muscular  action,  the  fluids  inconsequence 
passing  more  rapidly  to  the  distal  end  of  the  stomach. 


460  DU.aLISM    demonstrated. 

and  contiguous  parts,  this  cannot  readily  be  done  ;  hence  the 
ineffectual  efforts  of  the  gorged  leech  to  locomote,  for  the 
glutton  is  too  heavily  handicapped.  In  this  position,  how- 
ever (.F),  it  is  easy  to  perceive  that  the  dorsal  muscles  are 
greatly  expanded,  while  the  ventral  are  proportionately  con- 
tracted, in  order  to  produce  the  incurvated  condition  of  the 
body.  So,  then,  to  begin  with,  we  have  first,  the  general 
tody-expansion  (D),  which  must  be  maintained  for  retaining 
the  fluids.  In  the  second  place,  we  have  the  central  portions 
still  more  widely  expanded  for  receiving  the  fluids  in  the  ter- 
minal ends  {B).  Finally,  we  have  the  feet  expanded  and  the 
body  incurvated  by  expanding  the  dorsal  and  contracting  the 
ventral  muscles  (F1).  Hence,  there  can  be  no  earthly  doubt 
that  expansion  as  well  as  contraction  are  alike  subject  to  vol- 
untary control;  otherwise  it  were  utterly  impossible  to  pro- 
duce the-  above  phenomena.  The  amount  of  nervous  force 
absorbed  in  the  digestive  functions,  and  for  producing  circula- 
tion and  maintaining  equilibrium  in  pressure  in  the  condition 
of  repletion,  should  act  as  an  enormous  drain  upon  the  cen- 
tral nervous  system,  and  diminish  in  proportion  the  voluntary 
movements  ;  in  consequence,  he  would  become  inanimate  and 
sluggish  to  a  degree — a  condition  quite  common  to  gluttons  ; 
and  in  the  cold-blooded  animal,  in  which  respiration  and  cir- 
culation are  at  the  minimum,  of  course,  it  would  be  more 
marked,  inducing  prolonged  torpor. 

"We  now  pass  to  other  phenomena  in  which  expansile  action 
in  the  muscles  is  made  more  energetic,  notably  the  action  in 
the  tongue  of  the  frog. 

The  remarkable  rapidity  with  which  expansion  and  con- 
traction may  be  effected  in  the  muscles  has  forceful  illustra- 
tion in  the  tongue  of  the  frog.  Anatomy  :  The  tongue  of  the 
frog  is  conrposed  principally  of  two  muscles,  the  genio-glossus 
connected  with  the  mandible  (Fig.  190,  d,  a),  and  the  hyo- 
glossus  (c)  proceeding  from  the  posterior  cornua  of  the  hyoid 
bone,  together  with  the  cartilaginous  plate  (b)  which  is  pro- 
jected from  the  body  of  the  hyoid  bone  as  a  supporting  style, 
around  the  anterior  end  of  which  the  hyo-glossus  (c)  curves 
to  get  upon  the  dorsal  surface  ;  so  that  the  tongue  is  bent 
upon  itself,  the  terminal  end  presenting  toward  the   fauces. 


DUALISM     DEMONSTRATED.  461 

At  the  point  where  the  hyo-glossus  curves  over  the  end  of  the 
plate  it  forms  a  tendon,  which  facilitates  the  gliding  action, 
over  the  cartilage  when  the  organ  is  extended  and  retracted. 
When  forcibly  extended  in  the  dead  animal,  the  organ  reaches 
to  the  edge  of  the  mandible.  Such,  in  brief,  is  the  visible 
mechanics,  which  will  answer  the  purpose  of  description.. 
Now,  then,  the  animal  has  the  power  to  project  the  tongue  out 
of  the  mouth  beyond  the  mandible  with  the  rapidity  of  light- 
ning, extending  it  fully  two-thirds  the  length  of  the  body  for 
hooking  the  prey  and  compelling  it  into  the  mouth,  doing  it 
so  rapidly  that  the  eye  cannot  follow  it,  the  speed  in  extension 
and  retraction  being  so  great.  And  how  is  it  possible  to  explain 
this  phenomenon  by  the  monistic  theory  of  muscular  action  % 
Thus,  contraction  in  the  genio-glossus  (d,  a)  could  only  pull 
the  end  of  the  organ  to  the  edge  of  the  mandible  ;  and  with 


5 

c 

Fig.  190. — Diagrammatic  Representation  of  the  Tongue  of  a  Prog,  d,  Tongue,  reverted 
upon  the  cartilaginous  plate  (b)  of  the  hyoid  bone  ;  a,  d,  genio-glossus  ;  c,  hyo-glossus 
muscle. 

both  muscles  acting  simultaneously,  it  must  pull  the  reverted 
end  of  the  tongue  forward  upon  a  line  corresponding  with 
the  insertion  of  the  muscle  in  the  mandible,  or  straighten  it 
simply,  and,  of  course,  holding  it  in  this  position  during  the 
period  of  contraction,  which  is  all  this  action  in  the  muscle 
can  possibly  do,  but  which  only  puts  the  organ  in  a  position 
to  be  suddenly  projected  from  the  mouth, .  which  requires 
elongation  in  both  muscles  in  order  to  effect  it ;  otherwise  is 
utterly  impossible,  as  must  appear  obvious.  The  special  elec- 
trical current  for  producing  expansion  is  determined  into  the 
parts  under  the  volition  of  the  animal,  similar  to  what  occurs  in 
the  electrical  organ  of  fishes,  the  principle  being  the  same,  pro- 
jecting and  retracting  the  organ  with  the  rapidity  of  lightning 
by  reason  of  this  action  of  the  electrical  current ;  otherwise 
is  inexplicable.  Fortunately,  however,  we  have  a  means  of 
studying  the  action  in  the  muscles  ready  to  our  hand,  in 
which  the  movements  are  more  slowly  performed,  but  which. 


462  DUALISM     DEMONSTRATED. 

is  equally  conclusive  of  dualism  in  muscles  and  nervous  force, 
notably  in  the  tortoise,  to  which  attention  is  now  directed. 
Concerning  the  Movements  in  the  Head,  Neck  and  Tall  of  the 

Tortoise,  with  Reference  to  the  Principle  in  Musculation. 

Dualism  in  muscles  and  nervous  force  has  striking  illustration 
in  the  muscular  actions  for  producing  the  movements  in  the 
head,  neck  and  tail  of  the  tortoise  ;  but  in  order  to  place  the 
matter  fully  before  the  student,  it  will  be  necessary  to  make 
brief  reference  to  the  special  anatomy  in  the  parts  or  the  bones 
and  muscles  concerned  in  the  movements,  beginning  with  the 
cervical  vertebrae. 

Cervical  Vertebrae. — In  chelonia  the  cervical  vertebrae  are 
elongated,  eight  in  all,  the  last  one  (Fig.  191,  8),  articulating 
with  the  first  dorsal,  is  short  and  broad,  with  the  anterior 
surface  of  the  body  divided  into  two  transversely  elongated 
convexities,  the  posterior  forming  a  single  convex  surface, 
divided  into  two  lateral  facets,  corresponding  with  opposite 
depressions  and  elevations  in  the  first  dorsal,  forming  the 
cervico- dorsal  articulation  ;  the  fourth  (D)  is  elongated  and 
convex  at  both  ends,  fitting  into  shallow  depressions  in  the 
adjacent  vertebrae  so  as  to  admit  of  the  bending  and  folding 
up  of  the  cervical  chain  within  the  excavation,  the  whole 
forming  a  figure  very  similar  to  the  letter  S  reversed,  the 
upper  leg  at  the  carapace,  the  lower  (D)  at  the  plastron. 

Cervical  Muscles. — For  producing  the  movements  in  the 
cervical  vertebrae,  beginning  with  the  short  and  small  seg- 
mental muscles  in  immediate  contact  with  the  bones,  we  have 
first,  on  the  dorsal  region,  the  longus  colli  (t),  short  fasciculi 
extending  from  the  under  part  of  the  first  and  second  costal 
jxlates  and  first  dorsal  vertebra,  inclining  forward  and  inward, 
to  be  inserted  into  the  next  adjacent  vertebra,  rising  from  one 
to  be  inserted  in  the  other,  the  whole  chain  of  short  muscular 
links  thus  formed  receiving  this  name;  the  "  b  i  venter  cerm- 
cis,J  (e),  from  the  neural  plate  and  first  dorsal  to  be  inserted 
into  the  crest  and  fossae  of  the  occipital  bone,  and  from 
the  fifth  to  the  third  cervical  vertebrae,  to  be  inserted  into 
the  crests  of  the  occipital  and  temporal  bones  ;  upon  the 
sides  seven  short  fasciculi,  commencing  with  the  eighth  and 
ending  with  the  second  cervical  vertebrae,  known  as  "inter- 


DUALISM     DEMONSTRATED.  463 

transfer  sarii  colli,''''  with  other  fasciculi  more  obliquely  dis- 
posed, corresponding  with  the  fourth,  third,  second  and  first, 
known  as  the  "  intertr  answer  sarii  obliqiiV ;  upon  the  under 
surface  "scalenus"  proceeding  from  the  lower  three-fcurths  of 
the  scapula,  to  be  inserted  into  each  cervical  vertebra  from  the 
eighth  to  the  second  (s),  ' '  sternomastoicleus, ' '  proceeding  from 
the  sternum  to  be  inserted  into  the  mastoid  process ;  finally,  the 
powerful  uretra7iens  capitis  collique"  (M)  from  the  neural 
arches  and  spines  of  the  eighth  to  fifth  dorsal  vertebrae,  inclu- 
sive, to  be  inserted  by  four  tendinous  attachments,  the  long- 
est and  strongest  into  the  basi-occipital  fossa,  the  other  three 
into  the  fourth,  fifth  and  sixth  cervical  vertebrse ;  and  over 
all  of  them  the  great  cervical  muscular  sheath,  "  latissimus 
colli"  (Fig.  17:.',  21,  25),  inclosing  the  cervical  muscles  for 
retaining  them  in  position,  and  assisting  the  action  by  com- 
pelling the  articular  surfaces  to  glide  over  each  other ;  this 
will  serve  the  purpose  of  description.  And  it  will  not  be  neces- 
sary to  name  the  muscles  at  the  base  of  the  skull  connecting 
it  with  the  proximal  vertebra?,  since  they  could  do  no  work  in 
effecting  extension  and  retraction  of  the  organs  in  the  excava- 
tion The  same  applies  for  the  muscles  connected  with  the 
hyoidean  apparatus. 

Now,  then,  the  question,  How  are  the  head  and  neck  of 
the  tortoise  retracted  and  folded  up  in  the  excavation  (Fig. 
191),  thence  extended  again  (Fig.  192),  seeing  that  muscles 
act  between  the  points  of  origin  and  insertion  only?  And 
since  there  is  but  one  muscle — namely,  the  powerful  "retra- 
Tiens  capitis  collique''''  (M),  whose  points  of  origin  are  pos- 
terior to  the  cervical  flexure,  it  follows  that  this  is  the  mus- 
cle for  effecting  retraction ;  which  is  obvious  enough.  But 
the  animal  cannot  live  by  retracting  the  head  and  neck  ;  hence 
this  cannot  be  the  chief  function  of  this  muscle,  which  is  con- 
cerned in  extending  as  well  as  retracting  the  organs.  It  would 
scarcely  be  contended  that  the  small  fasciculi  connecting  the 
chain  of  bones  with  each  other  are  able  to  forcibly  pull  into 
extension  the  great  muscular  bundles  in  the  retrahens,  with 
all  the  advantage  of  leverage  against  them  in  addition.  ±say? 
it  cannot  be  entertained  for  a  single  moment,  even.  On  the 
contrary,  all  the  muscles  are  in  harmonious  concert  in  order  to 


4G4 


Dl'ALISM     DEMONSTRATED. 


effect  these  actions,  the  short  fasciculi  being  the  fine  adjust- 
ments for  compelling  the  articular  surfaces  to  glide  over  each 
other,  as  force  is  being  applied  by  the  powerful  muscles  for 
effecting  extension  and  retraction,  all  acting  together  and 
simultaneously,  the  muscular  sheath  at  the  same  time  facili- 
tating extension  and  retraction  by  contracting  and  expanding 
around  the  parts.  But  when  the  organs  are  being  extended,  the 
muscles  connected  with  the  sternum  and  scapulae  {scaleni, 
sternomastodei)  serve  to  guide  the  head  and  determine  it  in 


Fig.  191. — Skeleton  of  Tortoise  (Emys  Europcea)  (Owen),  showing  the  manner  in  which 
the  neck  and  head  are  folded  up  in  the  excavation,  forming  a  figure  similar  to  the 
letter  S  reversed,  the  upper  leg  (a)  at  the  carapace,  the  lower  (D)  at  the  plastron. 

the  direction  desired,  the  other  muscles  connecting  with  the 
vertebrae  and  head,  at  the  same  time  assisting.  And  the 
movements  not  only  involve  consentaneous  action  in  the 
muscles,  but  opposite  action  in  opposing  sets  of  muscles ; 
those  upon  the  dorsal  surface  of  the  vertebrae  undergoing 
elongation  as  those  upon  the  ventral  or  lower  aspect  are  un- 
dergoing contraction,  and  vice  versa  ;  otherwise,  extension  and 
retraction  of  the  parts  could  not  be  accomplished.  Finally, 
since  nervous  force  produces  hardness  in  the  muscles,  we  can 
readily  understand  how  the  great  muscular  bundles,  by  becom- 
ing more  and  more  rigid  as  they  elongate,  should  have  the  effect 
of  pushing  the  head  and  neck  out  of  the  excavation  for  effect- 
ing extension,  the  principle  being  the  same  as  in  the  worms, 


DUALISM    DEMONS  L  RAT  hD. 


465 


for  extending  and  erecting  the  soft  tubular  body  ;  one  set  of 
muscles  undergoing  elongation,  another  and  opposing  set  un- 
dergoing contraction,  both  sets  increasing  in  hardness  with 


S3 

qq' 

< 

rs 
4 

c+ 

& 
4 

p 

0 

Mi 

3 
b 

X 

0 

<1 
CD 
4 
eh 
CD 

c~ 

P 

B 

CD 

CD 

Oq 

eh 

tt 

5 

cr  c 

CD   w 
P     CD 
Pi    "i 
^    P 

PiCR 

0 

1 

0 

55 

05 

p" 

5' 

3' 

3' 

-~ 

3" 

CO 

5' 

B 

CD 
CD 
ft 

CD 
P 
P. 

4 
-1 
0 
B 

O 

pt 

o1 

S3 

CO 

; 

CD 

* 

X 

5" 

-. 

3 

72 

w 

eh 

a<3 
l-fc 

0 
4 

p 
Pi 

JS 

0} 

CD 

2., 

Oq 

0 

cr 

a>" 

1-+. 

5 
0 

w 

c 

CD 

0 

CD 

B 

P 

1 

3 
0 

Ma 

B 
eh 

P 
=1 
P< 

3' 

CO 

B- 

4 
eh 

0 

C5> 

4 

5' 

CO 
CD 

B 
Pi 

J' 

g 

on" 

3' 

£_ 

3 
p 
4 

3 

ca 
J' 

~. 

CO 

1 

CD 

eh 

eh 

B 

■* 

O' 

1 

f 

tr 
0 
eh 
p 

CB 

CO 
O 

tf 

3 
CO 

a 

eh 

O 

B, 

O 

B 

0 

>. 

CI 

0 

e-i. 

P 

••5 

CO 

0 

D1 
CD 

4 
Ha 

a 

CD 

B 
B 

09 

1 

a 

co 

5' 

P 
CO 

B 
4 

S 

•^ 

^ 

CO 

2 

p 

-T" 

CD 

4 

T 

^ 

s 

CO 

^ 

^ 

3 
a. 

K  3' 

W  iq 

3' 

Ml 
O 

p 

CD 

P 

5' 

CO 

4 

4 

c 

2 

CD 

3 

B 
P. 

CD 
O 

0 

TO 

s 

Co 

C5 

a 

B 

CD 

0 

0 
ft) 

CO 

co 

2 
B 

B 

CD 

M 

CO 

eh 

Bi 

4 
CO 

eh 

B 
Pi 

0 

Pi 

eh 
CD 

x 

0 

CD 

eh 

P 

P 

P, 
C 

c 

1. 

Ms 

- 

P" 

B 

-i 

< 

cr 

a 

CO 

P 

0" 

co" 

O 

^ 

— 

CO 

_P 

Ci 

c 

4 

CD 

CD 

< 

0 

CD 

4 
CO 

p 

CI. 

p- 

0 

Pi 
0 

i. 

r+- 

s- 

c 
0 
B 

0 

P 
0 

13 

B 

O 
CD 

4 

< 

Z 
B 

p. 

0 
0 

2 

4 

P 

-1 

CD 

1 

CD 

a" 

g 

3' 

CD 
4 

0" 

P^ 

P, 

eh 
O 
C 

p 

CD 

0 

0' 

CD 

3' 

eh 

r 

CD 

<4 

CD 
4 
el- 
CD 
& 
4 
P 

P^ 

CD 
CO 

P 

B 

-1 
CD 

eh 

P 

B 
P- 

Mi 

O 

B 
P 

5' 

Oq 

l-b 

3 

B 

CO 

2^ 

CD1 

B 

c 
H 

eh 

P 
& 

5' 
p 

0! 

B 
0 
0 

CD 
CO 
CO 

5" 

B 

pi 
Ba 

CO 

O 
CO 
X 

83 

0 

Ha 

CD 

B 
0 
CD 

W3 
O 
4 

O 

CD 

CD 

I 

XI 

2^ 

p' 
0 
CD 
B 

Pi 

0 
4 

'X 

P^ 

0 
O 

2 
►3' 

O 

eh 

B- 

4 

P 
<^ 

5' 

S1 

X 

Ms 

O 

ch 

CD 

iq 

4 

the  energy  in  the  movements.  It  follows  that  the  muscles  and 
nerves  possess  a  dual  function,  or  the  power  of  producing  ex- 
pansion and  contraction  under  the  volition  of  the  animal.  In 
the  tortoise  we  have  a  ready  means  of  demonstrating  this  cir- 


466  DUALISM     DEMONSTRATED. 

cumstance,  by  turning  the  animal  upon  its  back  simply,  then 
watching  the  result.  Remember,  it  may  be  the  first  time  it 
was  ever  upon  its  back,  so  that  it  would  have  to  bring  into 
action  special  muscles  for  the  first  time,  making  them  perform 
the  work  it  desires,  which  is  to  throw  itself  back  again  into  the 
normal  position,  or  upon  the  plastron.  Now,  then,  the  animal 
will  not  only  extend  the  head  and  neck  to  the  full  limit,  but 
curve  them  backward  over  the  edge  of  the  carapace,  stiff  and 
rod-like,  the  head  against  the  ground,  to  function  as  a  lever, 
which  is  used  vigorously  for  the  purpose ;  at  the  same  time, 
the  tail  also  is  extended  and  curved  backward  in  a  similar 
manner  over  the  carapace  ;  while  the  limbs  in  contact  with  the 
groundwork  vigorously,  catching  at  any  resisting  surface.  In 
this  manner  all  the  parts  are  employed,  till  at  last  the  work  is  ac- 


Fig.  193. — The  Deep  Muscles  in  the  Tail  of  the  Tortoise. — Bojanus. 

complished,  the  ground  favorable,  performed  quickly  and  deftly 
enough.  Pausing  to  inspect  the  muscles  of  the  tail,  we  find 
that  here  also  the  rjowerf ul  muscles  are  below  or  upon  the  ven- 
tral surface  (L94,  c  ;  193,  50,  5J,  52,  55),  the  short  fasciculi  upon 
the  lateral  aspect  being  removed  in  order  to  fully  expose  these 
muscles.  With  all  the  advantage  of  leverage  against  them,  it 
is  simply  impossible  for  the  short  fasciculi  to  pull  into  such 
forcible  extension  the  powerful  muscles  in  the  tail,  as  must 
appear  obvious. 

And  in  the  face  of  this  overwhelming  evidence  to  assert  that 
the  muscles  do  not  possess  dualism  (not  to  mention  the  law  in 
the  organs  of  circulation  compelling  this  circumstance,  in  order 
to  produce  the  rhythmical  changes  in  pressure),  that  they  can- 
not expand  as  well  as  contract,  exceeds  the  limits  in  reason  and 
fair  intelligence,  blindfold  though  he  be,  seeing  but  dimly 
through  the  veil  wrought  by  tradition  and  prejudice;  but 


DUALISM    DEMONSTRATED.  467 

easily  accounted  for  in  the  absence  of  the  law  underlying  the 
organism  and  the  fact  that  it  cannot  be  produced  artificially, 
tending  to  confirm  and  perpetuate  the  error.  But  we  now 
see  that  the  animal  itself  can  make  the  muscles  act  in  this 
manner  readily  enough,  nay,  must  do  so,  in  the  very  nature 
of  things ;  while  the  molecular  actions  it  involves,  admit 
of  distinct  mental  presentation,  and  are  natural  enough,  so 
that  recourse  to  the  supernatural  is  not  at  all  necessary 
any  more  than  for  the  phenomena  in  the  physical  sciences. 
But  the  evidence  may  be  still  more  simplified,  and  with  the 
object  of  bringing  out  in  clear  relief  this  fundamental  princi- 
ple in  animal  mechanics,  and  laying  bare  the  complicate 
arrangements  that  obtain  in  the  structures  and  organs  as  ad- 
justments with  the  special  functions,  we  again  have  recourse  to 
the  lower  animals,  this  time  the  conchif era,  which  will  subserve 
the  purpose  admirably. 

The  Action  in  Conchif  ers. — The  species  of  Mollusk  known 
as  conchifers,  or  bivalves,  also  lamellibranchiates,  from  the 
lamellar  branchiae  or  gills  (Fig.  194,  b,  g),  of  which  the  oyster 
and  scallop  are  the  most  common  examples,  have  all  the  soft 
parts  inclosed  within  two  concave  discs  (or  valves,  as  they  are 
commonly  called),  which  are  opened  and  closed  by  means  of  a 
hinge-joint,  and  powerful  muscles  (c)  connecting  with  the 
valves  immediately  in  front  of  the  joint,  which  is  situated  pos- 
teriorly ;  some  of  them  possess  two  such  muscles  widely  re- 
moved from  each  other,  which  induced  conchologists  to  form 
the  conchifera  into  two  sub-classes,  or  Monomyoria*  and 
Dimyoria,f  the  latter  the  more  numerous  class,  embracing  the 
mussel  tribe  and  many  others.  But  for  simplicity,  we  shall 
take  examples  from  the  first  class  only — namely,  scallop  and 
oyster.  It  is  the  common  impression,  born  of  the  monistic 
idea  of  muscular  function  (forced  to  it  from  this  circumstance), 
that  conchifera  open  the  valves  by  relaxing  the  muscles  sim- 
ply, closing  them  again  by  contracting  the  muscles,  the  resist- 
ance in  the  spring,  serving  to  open  them  when  the  muscles 
are  relaxed,  thereby  making  the  so-called  "  adductor  mus- 
cle" the  antagonist  of  the  wonderful  elastic  spring,  of  which 

*  j.iovo'i,  single  ;  ^vi,  a  muscle. 

jf  Si?,  twice  ;  mv?,  a  muscle — having  two  muscles. 


46d  DUALISM     DEMONSTRATED. 

we  have  heard  so  much ;  hence  the  name  to  this  muscle. 
]Sow,  then,  in  order  to  probe  this  matter  to  the  bottom,  I  made 
selection  of  the  oyster,  which  has  more  elastic  substance  in  the 
joint  than  any  other,  and  applying  a  trephine  over  the  inser- 
tion of  the  adductor  muscle,  succeeded  in  detaching  it.  "Well, 
the  valves  did  not  open,  but  remained  cloned.  Of  course, 
they  were  no  longer  firmly  held  together,  the  muscle  being  de- 
tached, but  the  valves  did  not  expand  as  they  should  have  done 
according  to  the  theory.  The  theory  broke  down  then  and  there. 
The  explanation  is  obvious.  Thus,  when  the  adductor  ex- 
pands, it  forces  the  valves  open  ;  reversing  the  action  contracts 
them  ;  while  the  special  arrangement  in  the  compressibly  elas- 
tic ligament  facilitates  the  action  simply,  making  it  more 
effective,  using  the  force  which  is  stored  in  it  during  contrac- 
tion for  assisting  expansion,  thereby  relieving  the  action  in  the 
adductor  to  this  extent.  And  that  this  is  the  principle  in  the 
mechanics,  is  fully  shown  by  the  following  circumstances, 
notably:  1.  The  valves  are  air-tight;  the  water  they  cor  tain 
will  not  run  out  of  them  when  inverted,  so  that  the  muscle 
would  have  to  expand  in  order  to  open  the  valves  ;  which  is 
announced  by  the  noise  they  make  when  fed,  the  inrush  of  air 
producing  the  characteristic  sound.  2.  The  action  in  the 
adductor  in  the  common  scallop  during  locomotion,  for  by 
energetically  expanding  and  contracting  the  adductor,  the 
animal  propels  itself  through  the  water,  effecting  a  retrogressive 
movement,  the  valves  widely  expanding  and  violently  beating 
the  water  by  contracting  against  it,  forcing  the  animal  back- 
ward through  the  water  by  this  means,  the  expansile  action 
removing  the  valves  beyond  the  point  where  they  could  be 
influenced  by  the  ligament.  The  wide  divarication  in  the 
valves  when  found  upon  the  beach,  the  soft  parts  all  out  of 
them,  is  due  to  the  drying  of  the  external  ligaments,  and  the 
shortening  this  produces  in  them  causes  the  valves  to  revert 
upon  the  beveled  edges,  so  that  when  fully  dried  the  valves 
are  nearly  at  right  angles.  But  nothing  of  the  kind  ever  takes 
place  when  the  ligaments  are  kept  moist,  as  in  the  living 
animal.  So,  then,  there  can  be  no  doubt  whatever  of  the 
existence  of  a  dual  force  in  the  adductor,  or  the  power  to  ex- 
pand and  contract  under  the  volition  of  the  animal.     And 


DUALISM     DEMONSTRATED. 


469 


turning  from  this,  again,  to  the  more  complex  movements  in 
the  higher  stages  in  development — notably,  the  action  in  the 
jaws — and  the  same  circumstance  is  readily  proven  on  these 
muscles  also,  the  stage  in  development,  for  obvious  reasons, 
making  no  difference  in  this  respect. 

The  Principle  in  Musculation  Applied  to  the  Action  in  the 
Jaws. — Briefly,  the  muscles  for  opening  and  closing  the  mouth 
are  the  masseter  (Fig.  195,  1)  and  temporalis  (Figs.  196,  2, 
195,  3),  and  internal  pterygoid  (Fig.  197),  one  for  each  side, 


Fig.  194. — Common  Scallop  (Pecten  Jacobcea). — Jones,  a,  h,  Lobes  of  the  mantle  ;  b,  g, 
branchial  laniellse  ;  I,  mouth  ;  k,  lips  ;  c,  adductor  muscle  ;  i,  "  foot"  ;  d,  e,f,  visceral 
mass,  principally  filled  with  ova  ;  n,  o,  convolutions  of  intestine,  seen  through  the 
transparent  tegumentary  membrane  ;  m,  anal  orifice. 

which  expand  and  contract  for  the  purpose,  the  principle 
being  the  same  as  in  the  so-called  "adductor"  in  bivalves; 
and  there  are  no  muscles  for  opening  the  mouth  by  con- 
tracting for  the  purpose,  barring  the  small  floating  muscles 
in  the  infra-maxillary  region  connected  with  the  hyoid  bone 
and  tongue  for  operating  these  organs  in  connection  with 
deglutition,  sucking,  speech,  etc.,  in  deglutition  serving  to 
approximate  the  base  of  the  tongue  against  the  epiglottis, 
so  as  to  close  the  glottis  and  prevent  intrusion  in  the  air- 
passages,  which  is  the  principal  function  of  the  genio- 
hyo-glossus,  genio-hyoid,  mylohyoid,  stylo-hyoid  and  digas- 
tric muscles  ;    the   genio-hyo-glossus,  in  sucking  by  curving 


470  DUALISM    DEMONSTRATED. 

the  tongue  in  the  longitudinal  axis  so  as  to  form  a  trough,  the 
mylo-hyoid  compressing  it  against  the  hard  palate,  forcing 
the  aliment  into  the  pharynx  in  deglutition,  the  other  muscles 
acting  in  concert,  the  jaws  closed  for  making  the  inferior 
maxilla  the  point  of  resistance.  In  other  words,  these  muscles 
relate  to  the  actions  in  the  tongue  and  hyoid  bone,  and  not  to 
the  jaws ;  therefore,  cannot  be  regarded  as  opponents  to  the 
massetal  and  temporal  muscles,  which  assist  their  action  by 
keeping  the  jaws  approximated  during  deglutition  and  suck- 
ing, granting,  for  sake  of  the  argument,  they  were  strong 
enough  to  antagonize  the  great  muscles  in  the  jaws,  which  can 
by  no  means  be  done.  But  to  remove  every  possible  excuse 
for  contrariety  of  opinion,  I  resolved  to  probe  this  matter  to 
the  bottom  also  ;  and  for  the  purpose  made  selection  of  a  Spitz, 
and,  placing  it  under  chloroform,  I  swept  the  scalpel  around 
the  base  of  the  lower  jaw  to  the  bone,  and,  twisting  the  bleed- 
ing vessels  with  torsion  forceps,  deflected  the  cutaneous  flap, 
together  with  the  platysma  myoides,  fully  exposing  the  infra- 
maxillary  regions,  dividing  first  the  anterior  portion  of  the 
digastric  near  the  insertion,  then  the  genio-hyoid  (dividing 
the  muscles  upon  both  sides),  the  mylo-hyoid  at  the  attach- 
ment to  the  hyoid  bone,  not  otherwise  interfering  with  the 
floor  of  the  mouth,  finally  snipping  some  of  the  fibres  of 
the  genio-hyo-glossus  connecting  with  the  hyoid  bone,  which 
completed  the  sections,  not  venturing  to  divide  the  muscle 
at  the  tubercles  for  fear  of  retraction  of  the  tongue,  pro- 
ducing suffocation  ;  nor  was  it  necessary,  since  these  fibres 
could  exert  no  influence  in  divaricating  the  jaws.  The  flap 
was  then  carefully  rejxlaced  and  secured  by  means  of  inter- 
rupted sutures.  And  the  animal  soon  returning  to  conscious- 
ness, I  punched  him  with  a  stick,  which  he  bit  viciously. 
That  settled  the  matter  there,  showing  conclusively  that  the 
masseters,  pterygoids  and  temporals  effect  the  movements  in 
the  jaws,  or  divaricate  and  close  them  by  means  of  their  action 
upon  the  inferior  maxilla,  which  is  done,  of  course,  by  means 
of  expansion  and  contraction.*    In  this  manner,    then,  the 

*  The  animal  took  no  nourishment  for  over  three  clays,  but  the  fourth  morn- 
ing the  milk  had  disappeared  from  the  basin,  he  having  drunk  it  some  time 
during  the  night ;  but  he  made  a  good  recovery,  the  wound  healing  rapidly. 


DUALISM     DEMOZSTSTKATED. 


471 


mechanics  in  the  jaws  are  laid  bare,  and  it  is  at  once  per- 
ceived that  the  principle  is  the  same  precisely  as  obtains 
in  bivalves,  through  the  action  in  the  so-called  "adductor 
muscle."  Thus,  the  masseters,  pterygoids  and  temporals  open 
the  mouth  by  expanding,  reversing  the  action  for  closing 
it,  the  orbicularis  oris  expanding  and  contracting  simulta- 
neously, all  the  parts  acting  in  concert,  while  coordination 
is  readily  effected  by  means  of  the  fifth  and  seventh  pairs  of 
nerves,  which  are  correlated  in  the  medulla  oblongata,  the 
fibres  being  also  distinctly  traced  into  it. 

The  great  relative  size  of  the  masseter  (Fig.  195,  1),  its  ad- 


Fig.  195. — Section  of  the  Upper  and  Lower  Jaw,  showing  position  and  voiume  ot  the 
masseter  muscle ;  reduced.— Bougery,  etc. 

vanced  position  in  front  of  the  hinge  (2),  closely  embracing  the 
angle  of  the  jaw  and  much  of  the  body  of  the  bone,  together 
with  the  number  and  disposition  of  the  muscular  bundles, 
stamp  it  at  once  as  the  chief  force  for  opening  and  closing  the 
jaws,  the  upper  one  being  fixed.  It  is  also  manifest  that  by 
contracting  the  bundles  connecting  with  the  malar  bone  the 
angle  and  ramus  of  the  jaw  would  have  the  effect  of  advancing 
the  jaw,  while  the  deep  portions  connected  with  the  zygomatic 
arch  and  the  body  of  the  bone  (not  shown  in  the  cut),  would 
have  the  opposite  effect,  or  retract  the  jaw  ;  thus  opening  and 
closing  the  jaw,  and  upon  occasion  moving  it  forward  and 
backward  for  triturating  the  food.     It  is  also  obvious  that  the 


472  DUALISM    DEMONSTRATED. 

position  of  the  muscles  is  most  advantageous  for  economizing 
force,  since  it  gives  the  greatest  leverage,  which  is  much  in  ex- 
cess of  the  temporal  (3)  inserted  in  the  coronoid  process, 
about  an  inch  only  in  front  of  the  hinge,  or  the  width  of  the 
ramus  itself  (Fig.  196,  1,  3).  The  obvious  purpose  of  this 
muscle  is  to  assist  the  masseter  in  opening  and  closing  the 
mouth  and  imparting  greater  force  to  the  bite,  being  well 
developed  in  carnivora.  Finally,  to  this  force  in  the  jaw  we 
must  add  the  action  in  the  pterygoids  (Fig.  197).  It  will  be 
seen  that  the  internal  pterygoid  is  also  a  powerful  muscle, 
resembling  the  masseter  in  form  and  the  direction  of  its 
fibres  ;  only  that  they  are  from  within  outward,  connecting 
the  bone  with  the  base  of  the  skull,  tending  to  pull  the 
bone  during  contraction  toward  the  median  line,  while  the 
action,  alternating  with  the  opposite  muscle,  rocks  it  from 
side  to  side  for  effecting  the  grinding  action  in  the  jaw.  The 
volume  of  the  muscle,  and  its  extensive  points  of  origin 
from  the  pterygoid  fossa,  the  pterygoid  plate  (external)  and 
palate  bone,  and  the  great  expanse  of  the  attachments  in 
the  ramus,  angle  and  body  of  the  jaw,  will  give  some  idea  of 
the  force  in  this  muscle.  The  pterygoids  expanding  simulta- 
neously would  exert  great  force  for  divaricating  the  jaws ; 
reversing  the  action  would  close  them  with  corresponding 
energy,  contracting  between  the  points  of  insertion  compelling 
this*  circumstance.  In  short,  the  pterygoids  and  masseters 
effect  the  grinding  action  in  the  jaws,  moving  the  lower  one 
upon  the  upper,  which  is  stationary  ;  while  the  temporals  in- 
crease the  action  for  divaricating  and  closing  the  jaws. 

Such,  in  brief,  is  the  tremendous  force  for  closing  the  jaws 
by  contraction,  while  there  are  no  opposing  muscles  for  open- 
ing it  by  contraction,  the  small  floating  muscles  in  the  infra- 
maxillary  region  already  referred  to,  with  the  hyoid  bone 
fixed  by  means  of  the  long,  slender  muscles  (sterno-hyoid, 
sterno-thyroid,  and  omo-hyoid)  connected  with  the  sternum  and 
scapulae,  may  have  some  influence  in  opening  the  mouth, 
but  that  they  are  not  important  muscles  in  this  respect,  is  now 
fully  proven  to  demonstration.  The  mouth  is  opened  with 
great  energy  and  celerity — opened  as  quickly  as  it  is  closed, 
and  with  considerable  force.     In  proof  of  this  latter  circum- 


DUALISM    DEMONSTRATED. 


473 


stance,  let  the  student  place  his  hand  under  the  jaw  with  the 
object  of  arresting  the  action,  and  he  will  be  at  once  convinced ; 
he  will  also  produce  pain  at  the  head  of  the  bone  from  strain 
to  the  ligaments,  produced  by  the  action  of  the  muscles  tend- 


Fig.  196. — Excision  of  the  Zygomatic  Arch,  showing  attachment  of  the  temporal  muscle 
(2) ;  in  the  coronoid  process  (1) ;  reduced.— Bougery,  ebc. 


ing  to  force  it  downward  and  backward,  the  anterior  portions 
being  immovable.  Furthermore,  he  will  find  by  palpating 
the  masseters  that  there  is  increasing  hardness,  or  the  same  as 
with  divarication  in  the  jaw,  and  when  fully  opened  the  mus- 
cles seem  as  hard  as  iron,  from  afflux  of  nervous  force  in  them  ; 
and  when  making  the   experiment  upon  himself,  a  painful 


474 


DUALISM     DEMONSTRATED. 


tension  in  the  joint  itself,  as  though  enduring  great  strain,  when 
the  mouth  is  opened  to  its  widest  extent. 

Finally,  this  action  in  the  muscles  would  explain  disloca- 
tion in  the  jaw  from  muscular  action,  and  the  special  arrange- 
ments that  obtain  in  the  parts  for  obviating  it,  otherwise 
inexplicable  ;  notably  the  existence  and  disposition  of  the 
external  pterygoid  muscles,  one  upon  either  side,  for  obviating 
displacement  ;  otherwise  inevitable.  For  example,  the  muscle 
extends  almost  horizontally  between  the  zygomatic  fossa  and 


Eig.  197. — The  Zygomatic  Arch  and  a  Portion  of  the  Ramus  of  the  Jaw  Removed, 
showing  position  of  the  pterygoid  muscles. — Gray. 

the  condyle  of  the  jaw  (Fig.  Iy7) — a  short,  thick,  powerful 
muscle,  connecting  the  condyle  with  the  superior  maxillary, 
palate,  and  sphenoid  bones,  and  spreading  out  widely  over 
them,  so  that  it  is  conical  in  shape,  the  small  end  being 
inserted  into  a  depression  in  front  of  the  neck  of  the  condyle, 
and  adjacent  part  of  the  inter -articular  fibro-cartilage,  the 
large  end  by  two  separate  and  broad  insertions  into  the  sphe- 
noid and  superior  maxillary  bones,  inclusive  of  the  palate 
bone,  so  that  an  effective  counter-force  applies  for  retaining 
the  head  of  the  bone  in  position  in  the  downward  and  back- 


DU.\LISM    DEMONSTRATED.  475 

ward  movements,  which  is  the  direction  of  the  force  in 
the  muscles  for  opening  the  month,  and  bnt  for  the  action  of 
the  external  pterygoids,  strain  and  dislocation  wonld  be  inevit- 
able in  every  widely  opened  month.  The  strong  counter-force, 
supplied  by  means  of  these  muscles,  applied  close  to  the 
articular  ends,  obviates  this.  Occasionally,  however,  the 
action  in  the  divaricating  muscles  is  excessive,  forcing  the 
body  of  the  bone  downward  and  backward  so  suddenly  and 
forcibly  as  to  dislocate  the  head,  throwing  it  out  of  the  articu- 
lating fossa,  the  action  in  the  external  pterygoid,  which  con- 
tracts simultaneously,  pulling  it  forward  over  the  crest  of  the 
fossa,  where  it  is  locked,  the  mouth  open  to  the  utmost  limit, 
produced  by  yawning. 

This  concentration  of  force  at  the  head  of  the  bone  from 
widely  diverging  points  in  the  base  of  the  skull,  effected  by 
the  two  heads  of  the  external  pterygoid,  corresponds  with 
the  plane  of  divarication  through  all  its  angles  as  the  counter- 
force  for  maintaining  the  head  of  the  bone  in  position ;  other- 
wise the  occasion  for  so  large  an  amount  of  muscular  force  in 
this  locality  is  inexplicable.  At  the  same  time,  the  insertion 
into  the  inter-articular  fibro-cartilage  would  tend  to  give  in- 
creased capacity  to  the  articular  cavity  for  effecting  extensive 
motion  in  the  head  of  the  bone  which  is  involved  in  the  grind- 
ing action,  showing  also  there  must  be  action  in  the  cartilage. 
In  this  manner,  then,  the  special  action  in  the  parts  is  readily 
explained,  while  the  adaptations  of  means  to  ends  would 
seem  to  be  absolutely  perfect. 

Passing  from  this  -as  being  sufficiently  animadverted  upon, 
we  next  take  up  the  action  in  the  circular  and  orbicular  mus- 
cles, to  note  how  readily  the  special  phenomena  appertaining 
to  them  are  also  readily  accounted  for  and  explained  by  the 
theory  of  a  dual  force  in  the  muscles ;  otherwise  are  inex- 
plicable. 

Concerning  the  Action  in  Orbicular  Muscles. — In  order  to 
illustrate  the  action  in  orbicular  muscles,  our  first  example 
shall  be  taken  from  the  iris,  for  the  reason  that  rude  force 
may  not  be  spoken  of  here,  where  everything  is  exquisitely 
organized  and  action  is  perfect ;  at  the  same  time,  there  is  a 
wide  arc  of  movement  in  the  muscles.     But  in  order  to  make 


476  Dl'ALISM     DEMONSTRATED. 

the  matter  fully  intelligible,  it  will  first  be  necessary  to  make 
a  brief  survey  of  the  special  anatomy  in  the  iris. 

Anatomy  of  the  iris :  Briefly,  the  iris  is  composed  of  two 
muscles,  the  circular  and  longitudinal,  the  muscular  layers  or 
lamellae,  and  a  delicate  stroma  of  fibrous  tissue,  in  which  the 
pigment  cells,  vessels  and  nerves  are  contained,  the  posterior 
surface  covered  by  a  thin  layer  of  connective  tissue  and  the 
uvea. 

*"The  nerves  of  the  iris  are  branches  of  the  ciliary  nerves 
of  the  choroid.  After  they  arrive  at  the  iris,  they  divide 
dichotomously  in  its  external  zone,  form  loops,  and  are  finally 
resolved  into  a  plexus  consisting  of  nerve-trunks  of  medium 
size.  In  this  plexus  may  be  remarked  an  interchange  of  the 
fibres  of  the  nerve-trunks,  thus  calling  to  mind  the  arrange- 
ment of  the  fibres  in  the  chiasma  nerv.  opticorum. 

"From  these  points  of  intersection  three  kinds  of  nerve- 
fibrils  take  their  origin  :  a,  pale  fibres,  in  all  probability  belong- 
ing to  the  sympathetica,  which  take  their  course  toward  the 
posterior  surface  of  the  iris  (consequently  toward  the  dilator), 
and  upon  it  form  an  exceedingly  fine  plexus  ;  b,  medullated 
fibres,  which  advance  to  the  anterior  surface,  and  these  are 
resolved  into  a  compact  network  of  fine  fibres — these  are  the 
sensitive  fibres  of  the  iris  ;  c,  finally,  a  third  plexus  is  dis- 
tributed within  the  sphincter  ;  its  delicate  fibres  are  for  the 
most  part  motor." 

"The  circular  muscle  (Fig.  19S,  a)  occupies  the  pupillary 
zone,  extending  outward  for  a  distance  of  0.09-1.3  mm.  It 
is  thinnest  at  the  pupillary  margin  .(0.10  mm.),  becomes 
thicker  externally,  and  near  the  outer  border  attains  a  thick 
nessof  0.25  mm."  According  to  ^Professor  Iwanhoff,  "the 
radiating  fibres  {bf  are  developed  from  the  bundles  of  the 
sphincter  as  an  uninterrupted  continuation  of  the  same.  Its 
beginning  is  formed  by  a  series  of  arched  interlacing  bundles, 
which  lie  partly  within  the  sphincter  (b,  a),  and  partly  on  its 
posterior  surface,  between  it  and  the  pigment  layer.     These 

*  A  Manual  of  Histology,  by  Prof.  S.  Strieker,  of  Vienna,  Austria,  in  co- 
operation with  Th.  Meynert,  F.  von  Recklinghausen,  MaxSchultze,W.Waldeyer 
and  others.  English  translation.  Art.  The  Organ  of  Vision.  II.  Tunica 
Vasculosa,  by  Prof.  A.  Iwanhoff,  p.  856. 


DUALISM    DEMONSTRATED.  477 

isolated  bundles,  after  they  have  passed  the  boundary  of  the 
sphincter,  unite  to  form  a  continuous  layer,  which  spreads 
over  the  entire  posterior  surface  of  the  iris ;  all  its  fibres  lie 
regularly  parallel  to  one  another,  and  all  are  arranged  in  lines 
radiating  from  the  pupillary  to  the  ciliary  margin.  .     ." 

The  dilator  pupillse  (Fig.  198,  b)  is  developed  from  the  bundles 
of  the  sphincter  as  an  uninterrupted  continuation  of  the 
same.  At  the  point  of  insertion  of  the  radiating  fibres  into 
the  ciliary  muscles  (Fig.  199,  b,  c)  they  bend  suddenly  upon 
themselves  and  return  to  the  pupillary  or  circular  muscle, 
some  of  the  loops  larger  than  others  and  lying  in  close  con- 
tact with  the  ciliary  muscles,  others  bending  higher  up  the 


Fig.  198. — Segment  of  the  Iris,  viewed  from  the  surface.—  Jcropheef.     a,  Sphincter  ;  bt 

dilator. 

central  portions  in  contact  only  (c,  c),  forming  themselves  into 
two  layers  (a,  a').  Italics  are  added.  From  this  it  transpires 
that  the  sphincter  is  the  chief  muscle  in  the  iris,  while  the 
radiating  fibres  serve  for  assisting  the  action  simply.  Indeed,, 
the  eminent  author  in  the  same  connection  uses  the  following- 
forcible  language :  ' '  The  literature  on  the  dilator  pupillse 
(radiating  fibres)  leads  us  unwillingly  to  the  belief  that  until 
the  time  of  Henle  the  existence  of  this  muscle  was  pre- 
supposed on  the  ground  of  absolute  physiological  necessity, 
rather  than  actually  demonstrated,"  r.ef erring  to  the  preva- 
lent belief  that  muscles  cannot  expand,  consequently  that 
there  must  be  opposing  muscles  for  pulling  the  sphincters 
into  extension,  but  which  we  positively  know  to  be  erroneous. 
Nay,  more  than  this,  one  portion  of  the  same  muscles  may 
expand  or  elongate  as  another  is  contracting  /id  shortening, 
both  taking  place  simultaneously ;    a  fact    which    is    fully 


478 


DUALISM     DEMONSTRATED. 


demonstrated  in  the  leech  during  imbibition  (Fig.  16) ;  indeed, 
is  seen  in  every  visible  movement  (Fig.  175).  The  former, 
however,  will  suffice.  Thus  at  the  oral  orifice  the  sphincter 
and  radiating  fibres  (which  answer  to  the  muscles  in  the  iris) 
expand  and  contract  together  and  simultaneously,  the  same 
applying  for  the  bulb-like  expansion  (1),  the  longitudi- 
nal and  circular  muscles  expanding  and  contracting  simul- 
taneously, in  the  part  adjacent  the  circular  fibres  are  con- 
tracted only,  in  the  next  adjacent  portion  (2)  both  sets  are 
again  expanded,  then  another  contraction  of  the  circular  less 
than  the  first ;  finally  both  sets  of  fibres  are  simultaneously 
expanded,  and  continue  to  expand  until  the  utmost  limit  is 
reached  (Fig.  17) ;  so  that  there  can  be  no  doubt  whatever 
that  both  actions  may  go  on  simultaneously  and  in  utmost 
concert. 


Fig.  199.— Arrangement  of  Muscular  Bundles  in  the  Iris.— 7b.  a,  a',  The  two  muscular 
layers  formed  by  the  radiating  fibres  ;  6,  b,  the  same  suddenly  bendirig  upon  them- 
selves before  attachment  to  the  ciliary  muscle  ;  c,  c,  same  fibres,  showing  degree  of 
approximation  to  the  ciliary  muscle  represented  in  subjacent  border  ground,  or  dark 
horizontal  shading. 

Making  the  same  deduction  upon  the  muscles  in  the  iris,  the 
special  phenomena  in  the  expansion  and  contraction  of  the 
pupil  are  at  once  made  intelligible,  the  two  sets  of  fibres,  circu- 
lar and  longitudinal,  expanding  and  contracting  together  and 
simultaneously  under  the  action  of  the  nervous  forces  in  the 
parts,  both  cooperating  to  this  end,  the  one  assisting  the  other. 
Thus,  for  expanding  the  pupil  the  portions  of  the  veil  con- 
tiguous to  the  sphincter  are  gently  pulled  aside  by  the  action 
in  the  radiating  fibres,  thereby  facilitating  the  action  in  the 


DUALISM     DEMONSTRATED.  479 

sphincter  ;  while,  for  contracting  the  pupil,  the  circular  fibres, 
by  pulling  upon  the  veil,  would  facilitate  the  action  in  the 
radiating  fibres,  so  that  the  action  is  reciprocal.  Nothing 
could  be  more  perfect  than  the  action  in  these  muscles,  but 
dualism  is  essential  to  it,  and  must  be  so  in  the  very  nature  of 
things,  as  in  no  other  way  could  a  balance  be  maintained.  And 
that  there  is  such  balance  in  nervous  force  to  the  muscles  is 
susceptible  of  demonstration.  For  example,  when  the  short 
root  from  the  third  pair  (motor  oculi)  to  the  ciliary  ganglion 
is  divided,  the  pupil  is  at  once  closed  ;  but  when  the  long  root 
from  the  ophthalmic  branch  of  the  fifth  pair  is  divided,  the 
action  is  the  very  opposite  ;  the  pupil  as  suddenly  expands  to 
the  utmost  limits,  which  shows  undoubtedly  opposite  character 
in  the  two  sets  of  nerve  fibres,  one  producing  contraction,  the 
other  expansion  ;  otherwise  is  inexplicable.  In  other  words, 
dualism  in  nervous  force. 

Then,  again,  either  condition  of  the  iris  involves  opposite 
action,  so  that  the  nerve-trunks  must  contain  nerve-fibres  to 
both  circular  and  radiating  muscles.  This  action  in  the  circu- 
lar muscles  brings  to  mind  the  action  upon  the  blood  vessels 
to  the  salivary  glands  (p.  284),  produced  by  irritation  of  the 
special  nerves,  stimulation  of  the  chorda  tympani  (Fig.  1  5,  c) 
causing  the  vascular  walls  to  expand,  producing  a  rush  of 
blood  into  them,  while  stimulation  of  the  sympathetic  fibres 
causing  them  to  contract,  thereby  diminishing  the  blood  in  pro- 
portion. In  the  one  case,  nervous  force  regulates  the  amount 
of  light  in  the  eye  ;  in  the  other,  the  amount  of  blood  in  the 
glands  ;  the  principle  the  same.  And  if  it  be,  as  alleged,  "  that 
the  radiating  fibres  are  an  uninterrupted  continuation  of  the 
circular,"  it  would  only  show,  as  in  the  case  of  the  leech  and 
worms,  that  a  given  portion  of  a  muscular  fibre  may  be  under- 
going expansion,  while  a  contiguous  portion  is  passing  through 
the  opposite  movement.  Moreover,  there  can  be  very  little 
doubt  but  that  the  undulations  passing  along  the  muscles 
when  stimulated,  involve  this  circumstance,  the  two  being 
inseparable  in  musculation.  Finally,  since  the  changes  of  form 
in  the  muscle-cells  involve  opposite  movements  in  the  molecular 
elements,  it  follows  that  the  nervous  apparatus  for  producing 
and  energizing  the  actions  should  be  in  correspondence  or  be 


4b0  DUALISM     DEMONSTRATED. 

also  dual,  which  would  include  the  power  of  coordinating 
them  in  the  functions,  whether  it  relate  to  circulation,  the  vol- 
untary movements,  or  the  action  in  the  iris,  etc.,  the  principle 
being  the  same  in  all.  And  since  it  depends  upon  opposite 
polarities  in  the  molecular  elements,  we  can  readily  understand 
why  electrical  force  should  precede  mechanical  change  in  the 
muscles.  All  this,  then,  is  at  once  made  intelligible  by  dualism 
in  muscles  and  nervous  force  ;  otherwise  is  utterly  inexplica- 
ble.   We  proceed  to  other  phenomena 

Concerning  the  Action  in  the  Oral  Muscles  — By  easy  men- 
tal process,  we  transfer  the  action  from  the  orbicular  and 
radiating  muscles  in  the  iris  to  the  orbicular  and  radiating 
muscles  in  the  mouth,  in  which  the  latter  fibres  are  gathered 
into  muscular  bundles  under  special  names,  in  place  of  being 
spread  out  in  a  uniform  layer  as  in  the  iris.  Indeed,  accord- 
ing to  some,  the  radiating  muscles  of  the  iris  in  the  white 
rabbit  (Fig.  2<)0,  b,  b)  are  gathered  into  bundles,  but  acting 
simultaneously  for  effecting  the  action  in  the  pupil,  whereas  in 
the  radiating  muscles  connecting  with  the  orbicularis  oris  (Fig. 
201,  7,  8,  9,  10,  11,  12,  13,  14)  the  action  may  be  confined  to 
several  muscles  at  a  time,  which  gives  the  power  of  express- 
ing the  moral  feelings  and  the  great  variety  of  movements 
characteristic  of  the  parts ;  at  the  same  time,  however,  it  in- 
volves the  same  principle  in  musculation  and  nervous  force 
for  producing  them,  related  muscles  expanding  and  contract- 
ing simultaneously  in  order  to  effect  the  actions  ;  otherwise 
impossible.  In  other  words,  the  orbicular  muscle  is  not  for- 
cibly pulled  into  extension  by  the  radiating  muscles  when  the 
mouth  is  opened,  but  expands  pari  passu  and  simultane- 
ously with  contraction  in  the  straight  muscles,  and  vice  versa 
when  the  mouth  is  closed,  all  the  parts  acting  in  concert  by 
means  of  the  special  nerves  for  producing  and  coordinating  the 
actions.  Furthermore,  this  is  in  correspondence  with  thev 
action  in  the  powerful  masseters,  etc.,  for  opening  and 
closing  the  maxillee,  already  referred  to,  and  which  involves 
corresponding  expansion  and  contraction  in  the  orbicu- 
laris oris ;  while  the  radiating  muscles,  by  pulling  upon  the 
labise  during  divarication,  facilitate  the  action  in  the  orbicu- 
laris, as  in  the  case  of  the  iris,  the  principle  being  the  same  ; 


DUALISM     DEMONSTRATED.  481 

while  by  means  of  the  fifth  and  seventh  pairs  of  nerves  run- 
ning into  every  portion,  the  actions  are  readily  produced  and 
coordinated.  It  is  all  plain  enough.  For  effecting  the  variety 
of  movements  in  the  parts,  it  calls  for  the  special  anatomical 
dispositions  that  obtain,  which  are  simply  perfect.  In  some 
animals  the  oral  slit  extends  very  far  back,  so  that  when  the 
jaws  are  widely  divaricated  it  reaches  almost  "from  ear  to 
ear."  It  is  the  case  in  carnivora,  and  when  the  lion  yawns  it 
sends  a  quiver  through  the  flesh.  But  critically  examine  that 
energetic  action  in  the  muscles  of  the  jaws  and  mouth,  and  the 
marvelous  concert  that  obtains  in  the  multitudinous  parts  is 
suitable  preparation  for  the  less  impressive  but  more  exten- 
sive movements  which  are  similarly  produced  and  coordinated 
in  the  medulla  oblongata,  or  the  respiratory,  circulatory  and 
locomotory  movements,  together  forming  a  connected  whole, 
as  has  already  been  fully  considered ;  so  that  there  can  be  no 
doubt  whatever  on  the  part  of  Nature  to  effect  these  ac- 
tions in  the  muscles  of  the  mouth  and  jaws.  For  example, 
when  the  jaws  are  divaricated  by  the  expansile  action  in 
the  masseters,  temporales  and  pterygoids,  the  orbicularis 
oris  expands  simultaneously  and  yari  passu  with  this  action  in 
these  muscles,  at  the  same  time  thezygomatici  (10, 1 1)  and  bucci- 
nator (IS)  contract  for  pulling  the  angles  of  the  mouth  toward 
the  masseters  (15),  the  whole  moving  together  and  simulta- 
neously the  same  as  in  the  muscles  of  the  iris  under  the  action 
of  the  nervous  apparatus,  which  produces  and  coordinates  the 
movements,  the  more  complicate  arrangements  in  the  muscles 
making  no  difference  in  this  respect  ;  while  for  closing  the 
mouth  the  action  is  simply  reversed,  the  utmost  concert  being 
maintained  in  the  parts  With  progressive  increase  of  expan- 
sile action  in  the  muscles  of  the  jaws,  there  is  corresponding 
increase  of  contractile  action  in  the  retractors  at  the  angles  of 
the  mouth,  so  that  the  overlying  integument  is  thrown  into  the 
characteristic  folds  in  front  of  the  masseters.  For  producing 
the  innumerable  local  actions  connected  with  the  lips,  the 
numerous  radiating  bundles  and  fasciculi  apply.  But  in  these 
cases  the  action  is  limited  to  the  special  localities,  in  which  a 
given  portion  of  the  orbicularis,  in  conjunction  with  the  mus- 
cle proceeding  from  it,  is  affected.     And  since  we  know  that 


482 


DUALISM    DEMONSTRATED. 


nervous  action  may  be  limited  to  a  part  of  a  muscle  only,  the 
comprehensive  arrangement  which  obtains  here  is  something 
beautiful  to  look  upon.  The  great  mobility  in  the  lips  is  un- 
doubtedly due  to  the  number  and  variety  of  the  local  actions, 
which  are  swiftly  changed  and  blended  upon  occasion  by  means 
of  the  nerves  for  operating  and  coordinating  the  structures. 
In  the  simple  act  of  nursing,  for  example,  it  requires  great 
mobility  in  the  lips  for  effecting  coaptation  with  the  nipple  and 
producing  the  movements  concerned  in  sucking ;  also,  as 
organs  of  prehension  in  taking  solid  food.  While  to  this, 
again,  must  be  added  the  functions  connected  with  oral  sounds, 


Fig.  200.— Muscular  Structure  of  the  Iris  of  a  White  Rabbit.— Carpenter,  a,  Sphincter 
of  the  pupil  ;  b,  6,  radiating  fasciculi  of  dilator  muscle  ;  c,  c,  connective  tissue  with 
its  corpuscles. 

and  as  channels  for  announcing  the  emotions.  Varied  as  are 
the  uses,  then,  the  numerous  adjustments  which  obtain  in  the 
lips  would  not  seem  excessive  ;  nor  the  grouping  of  the  radi- 
ating fibres  into  separate  muscular  bundles  be  deemed  strange, 
the  matter  being  altogether  different  from  the  simpler  action 
in  the  iris,  in  which  the  radiating  fibres  form  a  uniform  layer 
for  producing  the  action  in  the  screen  concerned  in  opening 
and  closing  the  pupil.  At  the  same  time,  it  is  equally  mani- 
fest that  the  same  mechanical  principle  applies  for  producing 
the  movements. 

We  pass  rapidly  over  this  anatomy,  the   object   being  to 
show  the  principle  in   the   mechanics   simply,    leaving    the 


DUALISM     DEMONSTRATED. 


483 


student  to  follow  it  at  his  leisure.  By  beginning  with  a  prin- 
ciple upon  which  to  base  the  mechanics,  and  proceeding  from 
the  simpler  to  the  more  complex  forms,  the  comprehensive 
arrangements  which  obtain  in  the  organs  are  seen  to  be  but  as 
many  beautiful  adaptations  of  means  to  ends,  while  order  and 
method  are  made  inevitable.  In  the  orbicularis  palpebrsB 
(4)  we  have  another  adaptation  of  the  mechanics  in  the  orbic- 


Pig.  201.  — Muscles  of  the  Head  and  Face. — Wilson  and  Buchanan.  1,  Frontal  portion 
of  the  occipito-frontalis  ;  2,  its  occipital  portion  ;  3,  its  aponeurosis  ;  4,  orbicularis 
palpebrarum,  which  conceals  the  corrugator  supercilii  and  tensor  tarsi ;  5,  pyra- 
midalis  nasi ;  6,  compressor  naris  ;  7,  orbicularis  oris  ;  8,  levator  labii  superioris 
alaeque  nasi — the  adjoining  fasciculus  between  ciphers  8  and  9  is  the  labial  portion  of 
the  muscle  ;  9,  levator  labii  superioris  proprius — the  lower  part  of  the  levator 
anguli  oris  is  seen  between  the  muscles  10  and  11  ;  10,  zygomaticus  minor  ;  11, 
zygomaticus  major  ;  12,  depressor  labii  inferioris  ;  13,  depressor  anguli  oris  ;  14, 
levator  labii  inferioris  ;  15,  superficial  portion  of  the  masseter  ;  16,  part  of  its  deep 
portion  ;  17,  attrahens  aurem  ;  18,  buccinator  ;  19,  attollens  aurem  ;  20,  temporal 
fascia  covering  the  temporal  muscle  ;  21,  retrahens  aurem  ;  22,  anterior  belly  of 
the  digastricus — the  tendon  is  seen  passing  through  its  aponeurotic  pulley  ;  23,  stylo- 
hyoid muscle  pierced  by  the  posterior  belly  of  the  digastricus  ;  24,  mylo-hyoideus  ; 
25,  upper  part  of  the  stern c-mastoid  ;  26,  upper  part  of  the  trapezius.  The  muscle 
between  25  and  26  is  the  splenius. 

ular  muscles.  By  reason  of  the  form  of  the  orbital  excava- 
tion, the  eye  occupying  the  cavity,  with  a  deep  depression 
round  it,  together  with  the  convexity  in  the  upper  lid,  which 
is  in  close  contact  with  the  ball,  a  special  arrangement  in  the 
muscles  for  opening  and  closing  the  eye  is  made  inevitable, 
as  the  ordinary  sphincter  would  not  answer.  Accordingly, 
we  have  comparatively  but  few  circular  fibres  in  the  upper 


484  dualism    di-:mo.\sti:ated. 

lid,  while  the  radiating  or  straight  fibres  form  a  broad  triangu- 
lar muscle  (levator  palpebral),  which  joins  the  tarsal  cartilage 
through  its  whole  length,  proceeding  thence  to  the  middle  line 
of  the  upper  portion  of  the  orbit,  to  be  attached  to  the  optic 
foramen  and  sheath  of  the  optic  nerve  at  the  very  bottom  of 
the  excavation.  Now,  then,  for  producing  the  wide  arc  of 
movement  in  the  upper  lid  is  the  explanation  for  the  great 
length  in  this  muscle,  since  it  is  the  provision  for  effecting 
extensive  contraction  and  expansion,  for  elevating  and  depress- 
ing the  lid,  while  the  breadth  of  the  attachment  in  the  tarsal 
cartilage  makes  the  action  simultaneous  over  the  whole  lid  ; 
at  the  same  time,  the  circular  fibres  serve  for  assisting  the 
action,  increasing  the  force  for  closing  the  lids,  and  by  ex- 
panding facilitate  retraction.  But  here,  as  elsewhere,  the 
two  sets  of  fibres  are  in  utmost  concert,  expanding  and  con- 
tracting simultaneously. 

And  so  we  might  proceed  from  sphincter  to  sphincter 
through  all  the  hollow  viscera,  in  illustration  of  the  antago- 
nism subsisting  between  the  circular  and  the  radiating  or  longi- 
tudinal muscles  and  the  principle  which  obtains  in  the 
mechanics,  but  further  extension  is  unnecessary,  for  it  is 
all  simple  enough,  and  we  cannot  be  misapprehended.  More- 
over, the  action  in  the  sphincters  has  already  been  sufficiently 
considered  in  connection  with  the  special  functions  in  the 
viscera,  in  which  it  was  shown  that  the  sphincters  expand  as 
the  sides  and  fundus  of  the  organ  contracts  and  vice  versa, 
the  principle  being  the  same  precisely  as  obtains  in  the  iris 
and  oral  muscles. 

Concerning  tlie  Action  in  Erectile  Tissue. — Finally,  we  come 
to  the  action  in  erectile  tissue,  in  which  we  have  a  different 
order  of  phenomena,  but  which  flow  out  of  the  properties  in  the 
muscles  and  in  nervous  force,  before  referred  to  ;  notably,  we 
have  enormous  expansion  with  hardness  in  the  structures, 
which,  after  a  short  duration,  again  pass  off,  leaving  them  in  a 
flaccid  and  shrunken  condition,  in  striking  contrast  to  what 
they  had  lately  been  when  erect  and  fully  expanded.  Further- 
more, the  action  is  rapidly  produced.  And  being  a  product  of 
nervous  force,  of  course,  it  is  susceptible  of  explanation  ;  but 
the  ineffectual  efforts  to  do  so  by  the  monistic  theory  of  mus- 


DUALISM   DEMONSTRATED.  485 

cular  and  nervous  action,  shows  necessity  for  introducing  a 
new  principle  in  animal  mechanics,  in  order  to  make  it  in- 
telligible. Physiologists,  in  seeking  an  explanation,  have 
recourse  to  the  action  in  the  penis,  in  which  it  was  thought  to 
be  due  to  obstructed  venous  return  from  the  organ  ;  but  that 
this  is  not  the  true  explanation  is  sufficiently  obvious  from  the 
action  in  the  tongue  of  the  chameleon,  where  it  cannot  apply, 
and  which  we  shall  now  briefly  consider. 

Action  in  the  Tongue  of  the  Chameleon. — Anatomy: 
Briefly,  the  organ  consists  of  two  sheaths,  an  internal  and  an 
external  sheath,  in  form  of  two  cylinders,  the  one  fitting  in 
the  other.  The  internal  is  formed  of  dense,  fibrous  and 
elastic  tissue,  the  fibres  decussating  at  right  angles  and  in- 
closing a  stile  or  supporting  cartilage,  projected  from  the 
body  of  the  hyoid  bone,  which  is  closely  invested  by  it,  es- 
pecially in  the  lower  portions  of  the  sheath,  where  the  attach- 
ment is  very  intimate  ;  but  in  the  anterior  portions  the  con- 
necting fibres  are  larger  and  longer,  and  admitting  of  con- 
siderable movement  of  the  membrane  upon  the  stile,  permitting 
the  organ  in  the  relaxed  condition  to  be  folded  up  upon  it 
much  "like  the  seam  of  a  dress  upon  a  bodkin"  (C,  c),  the  end 
of  the  stile  presenting  in  the  lower  lip  of  the  bulb  (A,  a).  And 
when  thus  folded  within  the  mouth,  the  organ  measures  about 
one  and  one-half  inches;  but  when  extended,  however,  it  is  from 
six  to  seven  inches  in  length,  and  projecting  far  beyond  the  end 
of  the  stile  (B,  b).  But  there  is  no  canal,  the  elastic  tissue 
fibres  filling  up  and  occupying  the  portion  corresponding  with 
the  canal,  the  stile  making  its  way  through  the  interlacements 
when  the  organ  is  being  folded  up  upon  it.  The  large  bulb 
forming  the  head  of  the  organ  is  furnished  with  a  short  upper 
and  a  long  lower  lip  (B,  c,  b),  which  are  approximated  in  the 
relaxed  condition,  so  as  to  form  a  transverse  fissure,  but  in 
erection  they  are  expanded,  presenting  a  funnel-shaped  open- 
ing (Fig.  203),  with  the  lower  lip  extended  and  curving  a  little 
upward.  The  opening  is  also  supplied  with  an  orbicular  mus- 
cle, which  would  explain  the  opening  and  closing  of  the  lips 
for  seizing  the  insect,  while  the  glutinous  secretions  for  lubri- 
cating the  parts  would  cause  it  to  adhere.  Finally,  a  thin, 
muscular  layer  (hyo-glossus),  composed  of  longitudinal  fibres, 


4fr6 


DUALISM   DEMONSTKATED. 


extends  the  whole  length  of  the  organ  upon  the  lateral  aspects 
(one  on  either  side)  ;  while  a  large  nerve  courses  on  top  of  it  to 
reach  the  head  of  the  organ. 

The  sheath  corresponding  with  the  mucous  membrane  is 
formed  of  white  fibrous  tissue,  the  fibres  also  decussating  at  va- 
rious angles,  some  running  transversely,  others  longitudinally. 
(There  is  a  difference  of  opinion  in  regard  to  the  histological 
character  of  some  of  these  fibres,  which  are  believed,  on  the 

IN 


Fig.  202. — Tongue  of  Chameleon. — Cyclopaedia  of  Anatomy  ;  Art.  Tongue.  H.  Hyde 
Salter.  A,  retracted  ;  B,  elongated  ;  C,  seen  from  beneath,  in  situ,  by  incision  and 
separation  of  the  integument  ;  a,  anterior  portion,  dome  shaped  ;  b,  posterior  por 
tion  :  c,  upper  lip  of  tongue  ;  b,  lower  lip  of  tongue  ;  e,  genio-hyoid  muscle  ;  /,  cerato- 
maxillary  muscle  ;  g,  sterno-hyoid  muscle  ;  h,  cerato-sternal  muscle  ;  i,  omo-hyoid 
muscle  ;  k,  apex  of  greater  cornua  of  hyoid  bone. 

part  of  some,  to  be  unstriped  muscle  elements ;  but  since 
nervous  force  is  not  limited  in  its  action  to  the  muscles,  it 
need  not  concern  us.)  Thus  constituted,  the  organ  is  made  to 
ride  to  and  fro,  forward  and  backward,  by  the  action  in  the 
genio-hyoid  (A,  e)  and  omo-hyoid  (g)  muscles,  while  it  is 
raised  or  lowered  by  means  of  guy-muscles  (h,f)  attached  to 
the  cornua  (Jc)  of  the  hyoid  bone  for  aiming  it  at  the  object 
— much  in  the  manner  of  a  cannon  upon  the  carriage.  For 
effecting  the  movements  in  the  opposing  sets  of  muscles,  of 
course,  it  requires  dual  action,  the  one  expanding  as  the  other 


DUALISM   DEMONSTRATED.  487 

is  contracting,  and  vice  versa ;  in  this  manner  materially 
assisting  each  other.  1  hus,  as  the  genio-hyoid  (A,  e)  contracts 
for  pulling  the  hyoid  carriage  toward  the  oral  orifice,  the  omo- 
hyoid (g)  elongates  for  pushing  it  along,  and  vice  versa  for  run- 
ning it  back  after  erection.  The  same  remarks  apply  for  the 
guy-muscles,  which  are  also  made  tense  in  this  manner  for  regu- 
lating the  point  of  elevation.  Finally,  when  projected,  it 
leaps  off  the  stile  in  energetic  action,  going  straight  to  the 
object  as  though  shot  ont  of  a  pistol,  and,  striking  the  target, 
is  as  rapidly  returned  upon  the  stile  in  the  folded  condition,  as 
previously.  For  effecting  the  former  action,  the  circular  fibres 
are  suddenly  contracted  upon  the  network  of  elastic  fibres  oc- 
cupying the  canal,  while  the  longitudinal  fibres  elongate,  and 
the  hardening  this  produces  pushes  it  off  the  stile  in  the  erect 
position,  and  serving  to  hold  it,  rod-Wee,  from  the  end  of  the 
supporting  staff  (20-J) ;  otherwise  it  would  fall  and  the  end 
of  the  staff  be  caught  in  the  fibres.  Again,  there  is  no  central 
canal,  for  the  circular  fibres  contract  upon  the  network  so  as 
to  convert  the  whole  into  a  solid,  stiff  mass,  and  greatly  re- 
duced in  size  in  consequence  (Fig.  20H),  with  the  head  con- 
tracted and  elongated ;  while  for  effecting  retraction  the  action 
is  simply  reversed,  the  longitudinal  fibres  contracting  and  the 
circular  expanding,  and  which  would  include,  of  course,  the 
central  network,  for  nervous  force  pervades  the  whole  of  it ; 
while  the  hardening  this  produces  in  the  structures,  as  in  ex- 
ample of  the  soft  body  of  the  worms,  enables  the  actions  to  be 
produced ;  otherwise  is  inexplicable.  The  tremulous  action . 
in  the  organ  previous  to  launching  is  produced  by  the  guy- 
muscles  (B,  h,  f)fov  aiming  the  organ,  while  the  bolt  itself  is 
suggestive  of  the  discharge  of  electric  currents  through  the 
organ  by  means  of  the  large  lateral  nerves.  But  the  partial 
and  more  deliberate  protrusion  which  precedes  the  final  dis- 
charge is  due  to  the  action  of  the  muscles  on  the  hyoid  bone, 
or  the  genio-hyoid  and  omo-hyoid  muscles  ;  but  with  the  re- 
mainder of  the  phenomenon  they  can  have  but  little  to  do, 
save  for  running  the  carriage  back  again  at  the  end  of  the  per- 
formance to  the  place  it  occupies  in  a  state  of  rest.  Finally, 
it  should  be  borne  in  mind,  in  this  connection,  that  when  the 
organ  is  thus  projected  to  the  extremest  limit  in  extension,  it 


488  DUALISM  DEMONSTRATED. 

still  retains  perfect  control  of  all  its  parts,  else  the  movements 
in  the  end  of  the  organ  conld  not  be  produced  for  seizing  the 
insect,  nor  the  sheaths  be  returned  and  folded  up  upon  the 
stile  ;  so  that  in  no  sense  can  the  action  be  regarded  as  a  pas- 
sive one,  or  the  product  of  a  single  force  simply  ;  on  the 
contrary,  it  is  product  of  a  dual  force  subject  to  the  volition  of 
the  animal,  and  involving  perfect  coordination  in  all  the  parts 
for  effecting  it. 

Concerning  the  Action  in  the  Penis. — This  power  in  nervous 
force  for  producing  expansion  in  the  muscles,  elastic  and 
fibrous  tissues  would  explain  the   phenomena  in  the  penis, 


Pig.  203. — Tongue  of  Chameleon,  when  in  action. — Jones. 

hitherto  inexplicable,  making  this  also  plain  and  easily  under- 
stood, the  means  to  ends  being  very  perfect. 

Anatomy :  Briefly,  the  penile  organ  is  composed  of  a  mass 
of  erectile  tissue,  in  form  of  three  cylindrical  compartments, 
known  as  corpora  cavernosa  (which  occupy  the  upper  surface) 
and  corpus  spongiosum  (containing  the  urethra),  the  lower 
surface,  occupying  the  central  groove  formed  by  the  corpora 
cavernosa,  expanding  over  the  ends  to  form  the  glans  penis, 
which  is  firmly  secured  in  position  by  means  of  connective- 
tissue  fibres.  The  several  bodies  are  contained  in  a  separate 
sheath,  com  posed  of  elastic  and  fibrous  tissue  similar  to  the 
elastic  coat  in  the  arteries  (Fig.  204,  h),  dense  and  thick,  and 
closely  bound  together  from  the  intermingling  of  the  fibres 
along  the  sides,  while  within  this  outer  tunic  is  a  muscular 
cylinder,  the  two  intimately  interblending  to  form  the  com- 
mon wall  (h).  From  the  interior  of  this  sheath  (tunica 
albuginia)  proceed  numerous  bands  and  cords  (trabecules), 
which  intersect,  dividing  and  subdividing  the  interior  into 


DUALISM   DEMONSTRATED. 


489 


numerous  separate  compartments  (d,  d),  lined  by  laminated 
epithelium — a  continuation,  in  short,  of  the  endothelial  lining 
of  the  veins,  with  which  they  freely  communicate  ;  and  so  free 
and  intimate  is  this  communication  with  the  veins  that  they 
were  at  first  taken  to  be  mere  venous  dilatations,  but  are  vascular 
spaces  simply,  formed  by  the  trabeculse,  into  which  the  endo- 
thelial lining  of  the  vessels  is  projected,  or  similar  to  what 


Fig.  204. — Transverse  Section  through  the  Spongy  Portion  of  the  Urethra  (corpus 
spongiosum  urethras). — Klein,  a, '  Epithelium  ;  6,  tunica  mucosa;  c,  muscular 
cords  ;  d,  vascular  spaces  of  the  corpus  cavernosum  ;  e,  glands  ;  /,  excretory  duct 
of  gland  ;  g,  longitudinal  muscles  ;  h,  tunica  albuginia. 

occurs  in  the  uterine  and  placental  sinuses  (Fig.  137,  c,  c). 
Finally,  we  have  large  arterial  capillaries  emptying  into  these 
vascular  spaces  by  funnel-shaped  openings,  for  producing  rapid 
afflux  of  blood  in  the  parts,  and  embossed  by  dense  nervous 
plexuses  connecting  with  the  nerves  and  structures  in  the 
penis,  on  the  one  hand,  and  with  the  hypogastric  plexus  on  the 
other,  by  means  of  which  the  expansile  action  in  the  organ  and 
the  requisite  blood-supply  for  filling  and  distending  the  vascu- 
lar spaces  during  the  sexual  orgasm  are  readily  produced; 


490  DUALISM   DEMONSTRATED. 

the  blood  serving  to  give  volume  to  the  organ ;  otherwise  it 
would  be  needle-like  ;  hence  these  vascular  arrangements 

This  will  serve  the  purpose  of  description.  In  the  flaccid 
condition  the  organ  is  comparatively  small,  the  dense  fibrous 
and  elastic  sheath  and  muscular  trabecule  contracted,  the 
organ  less  than  one-third  the  size  in  the  state  of  erection. 
Now,  then,  the  question :  How  is  this  wonderful  transforma- 
tion effected?  Well,  the  only  explanation  that  present  physi- 
ology can  offer  (forced  to  it  by  the  monistic  theory  of  muscu- 
lar and  nervous  force)  is  the  one  of  venous  obstruction  produced 
by  contraction  of  the  muscles  at  the  root  of  the  organ,  com- 
pressing the  veins,  thereby  causing  the  blood  to  dam  back  in 
the  vascular  spaces  ;  this,  together  with  similar  obstruction  at 
the  venous  outlets  of  the  vascular  spaces  produced  by  disten- 
sion (as  suggested  by  others,  not  feeling  sure  that  the  former 
would  be  sufficient).  But  the  following  incontrovertible  facts 
show  conclusively  that  neither  is  correct,  nor  can  it  be  in  the 
very  nature  of  things.     Notably  : 

a.  The  walls  of  the  veins,  in  common  with  the  universal 
rule,  are  very  distensible,  yielding  readily  under  pres- 
sure ;  hence,  it  were  utterly  impossible  to  force  open  the 
sinuses  by  this  means  or  choking  the  veins,  since  it  would 
involve  the  forcible  distension  and  pulling  of  the  thick 
trabecule  and  the  dense  fibrous  sheaths  inclosing  the  organ 
to  fully  double  their  length  (not  to  mention  the  extraor- 
dinary hardness  which  accompanies  it,  and  which,  of  course, 
would  require  corresponding  force  for  producing  it,  involv- 
ing a  degree  of  strain  to  the  vessels,  painful  even  to  contem- 
plate), the  veins  as  the  point  of  resistance,  the  absurdity  of 
the  proposition  is  too  obvious  for  controversy.  The  veins 
would  burst  under  the  force.  Hence,  it  cannot  be  entertained 
for  a  single  moment  even. 

b.  It  would  involve  arrest  of  circulation  in  the  parts,  which 
cannot  be  thought  of,  for  it  would  at  once  put  the  organ  in  peril, 
since  this  is  simply  strangulation — can  be  nothing  else  ;  hence, 
in  prolonged  priapism,  incidental  to  specific  disease  and  abnor- 
mal irritati<  >n,  would  inevitably  produce  necrosis  or  death  of  the 
organ,  which  never  takes  place  from  this  cause.  Moreover,  it 
has  been  demonstrated  (Eckhard)  that  in  place  of  arrest,  or 


DUALISM  DEMONSTKATED.  491 

even  partial  arrest,  circulation  is  vastly  increased  in  the  organ 
during  erection,  the  arteries  being  greatly  expanded  and  the 
veins  discharging  an  unusual  amount  of  blood. 

c.  Pressure  in  the  sinuses  could  not  exceed  pressure  in 
the  arterial  system,  else  reflux  should  inevitably  occur,  the 
blood  escaping  in  this  direction,  and  so  relieving  the  struc- 
tures ;  and  since  pressure  in  the  sinuses  could  not  exceed 
pressure  in  the  arterial  system,  this  would  not  be  competent 
to  force  open  the  fibrous  sheaths  and  trabeculse,  and  produce 
the  characteristic  hardness  ;  granting,  for  sake  of  the  argu- 
ment, the  veins  could  offer  sufficient  resistance,  which  can  by 
no  means  be  done.  Hence  the  phenomena  could  not  be  pro- 
duced by  damming  the  blood  in  the  sinuses. 

d.  It  would  not  account  for  decline  of  erectile  power  with 
advancing  years  and  the  waning  of  general  muscular  force 
(they  go  together),  for  circulation  is  unimpaired  in  the  parts, 
and  secretion  is  abundant.  Nevertheless,  erectile  power  in 
the  organ  corresponds  with  the  waxiug  and  waning  of  muscu- 
lar force,  the  minimum  in  extreme  age. 

e.  It  would  not  explain  the  sudden  flaccidity  and  collapse 
of  the  organ  when  in  the  state  of  erection  from  mental  causes, 
disgust,  fear,  etc.,  since  it  would  be  utterly  impossible  for  the 
blood  to  escape  so  rapidly  in  order  to  effect  it. 

f.  Last,  but  not  least,  it  is  utterly  incompetent  to  explain  the 
anatomy  in  the  organ,  notably  the  number  of  the  muscles  tra- 
versing the  erectile  tissue  and  forming  the  walls  of  the  sinuses, 
together  with  a  muscular  cylinder  far  inclosing  them  beneath 
the  containing  sheath  of  fibrous  and  elastic  tissue.  Nor  why 
the  arterial  capillaries  should  terminate  in  the  sinuses  by 
funnel-shaped  openings. 

The  explanation  is  easy,  embracing  all  the  phenomena,  as  a 
matter  of  course  :  notably,  1.  The  containing  sheath  and  tra- 
becule expand  under  afflux  of  nervous  force  in  the  parts,  the 
amount  of  this  determining  the  degree  in  hardness.  2. 
Simultaneous  with  this  expansile  action  in  the  erectile  tissue, 
the  arterial  capillaries  and  trunks  expand  correspondingly  for 
filing  the  sinuses  or  vascular  interspaces,  the  suction  force  in 
the  latter,  together  with  the  pressure  in  the  arterial  system, 
compelling  them  to  be  filled  instantaneously,  thereby  pro- 


492  DUALISM   DEMONSTRATED. 

during  enormous  afflux  of  blood  in  the  sinuses,  the  veins 
serving  to  carry  off  the  excess,  so  as  to  produce  a  current 
through  the  organ  and  prevent  reflux  into  the  arterial  sys- 
tem ;  at  the  same  time  maintaining  a  flow  of  arterial  blood 
through  the  structures  for  evolving  the  force  in  the  muscles, 
which  is  essential  to  their  action,  in  this  manner  obviating 
strain  and  preserving  vitality,  and  which  cannot  be  done  in 
any  other  way ;  while  all  the  circumstances  in  the  anatomy  of 
the  organ,  inclusive  of  the  funnel-shaped  openings  of  the 
arterial  capillaries,  fall  readily  into  line  at  the  proper  time  and 
place,  leaving  no  outstanding  quantity  refusing  absorption, 
thereby  proving  the  correctness  of  the  premises.  It  is  need- 
less to  extend  the  matter. 

From  this  we  proceed  to  other  portions  of  the  genital  appa- 
ratus, making  a  brief  survey  of  them  also,  for  completing  the 
description  so  far  as  it  appertains  to  the  special  mechanics 
which  we  wish  to  elucidate.     It  will  not  detain  us. 

Concerning  the  Action  in  the  Vas  Deferens  and  the  Prin- 
ciple it  Involves. — There  is  no  hollow  organ  in  the  body  more 
muscular,  comparatively,  than  the  vas  deferens  (Fig.  205),  not 
excepting  the  heart  itself,  the  cavitary  space  (Fig.  39)  being 
relatively  larger,  comparing  part  with  part.  Thus,  the  vas 
deferens  has  three  great  muscular  layers,  the  inner  (c),  middle 
(d)  and  outer  layer  (e)  ;  the  inner  and  outer  layers  composed 
of  longitudinal,  the  middle  of  circular  fibres,  the  muscles  being 
thus  systematically  arranged.  Now,  then,  since  muscles 
relate  to  work,  why  all  this  force  for  the  vas.  deferens? 
This  also  is  easily  answered  ;  notably,  the  spermatic  fluid  is 
highly  albuminous  and  very  tenacious  ;  moreover,  has  to  ascend 
the  tube  perpendicularly  (Fig.  206) ;  hence,  this  degree  of  force 
for  compelling  circulation.  But  how  compel  %  By  contracting 
from  below  up  !  Very  well.  But  this  involves  several  impor- 
tant circumstances  for  which  we  have  been  contending,  nota- 
bly :  1.  The  fact  of  "automatism  in  the  organs,"  which  is 
absolutely  essential  to  the  performance  of  their  functions  ;  also 
involving  local  mind  centres  for  coordinating  the  movements, 
or  the  same  as  obtains  for  the  separate  independent  organism, 
living  and  sustaining  existence  by  itself.  2.  It  involves  the 
principle  for  producing  movements  in  the  contents  by  means 


ACTION   IX   THE   YAS   DEFERENS. 


^93 


Kg.  205.— Transverse  Section  through  the  Commencing  Portion  of  the  Vas  Deferens. — 
Klein,  n,  Epithelium  ;  b,  tunica  mucosa  ;  c,  inner  ;  d,  middle  ;  e,  outer  muscular- 
layer  ;  /,  bundles  of  the  cremaster  internus  ;  g,  vein  containing  muscles  in  its  walls. 


Tunica  Viyinalij 
Tunica  JMluj  in  ia 


Klg.  a06. — Vertical  Section  of  the  Testicle,  to  show  the  arrangement  of  the  ducts. — Gray. 


19  i  DUALISM     IN    NERVOUS   FORCE. 

of  "rhythmical  changes  in  pressure";  therefore,  is  based  upon 
pressure,  which  is  fundamental  in  the  organism  for  compelling 
circulation  in  the  measure  of  the  physiological  requirements, 
this  whether  it  relate  to  the  blood  itself,  the  juices  in  the  struc- 
tures, or  the  secretions  in  the  organs. 

3.  It  involves  dualism  in  muscles  and  nervous  force  in 
order  to  effect  the  actions,  or  the  power  to  produce  expansion 
and  low  pressure  in  one  portion,  with  simultaneous  contraction 
and  high  pressure  in  a  contiguous  portion,  whereby  rapid 
movement  in  the  contents,  with  obviation  of  friction  and 
strain,  are  readily  effected.  A  most  notable  circumstance  in 
the  vas  deferens  is  the  absence  of  valves  for  sustaining 
the  fluids  and  obviating  reflux ;  this,  notwithstanding  the 
ascent  is  perpendicular,  in  order  to  reach  the  vesiculse  semi- 
nales  or  seminal  reservoirs  at  the  base  of  the  bladder  (Fig.  *207), 
where  the  fluid  is  stored  for  favorable  opportunity.  But  the 
mucous  membrane  is  in  longitudinal  folds,  that  increase  in  num- 
ber in  the  ampulla,  the  dilated  lower  extremity  (Fig.  206),  in 
which  the  muscles  are  also  increased,  and  which,  of  course,  have 
relation  to  and  correspond  with  the  amount  of  work  that  is  in- 
volved for  lifting  the  fluid  up  the  tube.  In  other  words,  the  ar- 
rangements which  obtain  are  similar  to  what  occurs  in  the  bron- 
chial tubes,  cesophagus,  and  stomach,  for  compelling  movement 
in  the  contents,  the  folds  in  the  mucous  membrane,  as  in  the 
other  cases,  being  necessary  adjustments  with  the  action  in  the 
muscles,  permitting  them  to  expand  and  contract  upon  the 
contents  in  the  measure  of  the  requirements.  A  circumstance 
which  strengthens  this  conclusion  is  the  extension  of  the  mus- 
cles into  the  mucous  membrane,  the  fibres  traversing  it  to  the 
epithelium,  so  that  simultaneous  action  in  all  the  parts  is 
thereby  made  inevitable.  If  will  not  be  necessary  to  the  argu- 
ment to  enter  into  the  minutia  of  the  beautiful  and  won- 
derful anatomy  in  the  testis,  the  number,  variety,  and  ex- 
tent of  the  tubes  (the  totality  said  to  approximate  a  mile 
in  length);  suffice  it  to  say,  that  we  have  this  delicate  struct- 
ure, the  problem  being  to  remove  the  secretions  as  fast  as 
they  are  formed.  And  it  is  not  to  be  supposed,  for  a  single 
moment  even,  that  the  powerful  ampulla,  into  which  they  are 
previously  discharged  (passing  through  a  narrow  muscular 


DT.ALISM    IN    NERVOUS   FORCE.  495 

portion  to  reach  it),  is  forcibly  distended  by  the  tnbules  by 
compelling  the  fluid  into  it ;  but,  on  the  contrary,  that  this 
expands  from  the  sensory  impressions  in  the  mucous  surface 
produced  by  the  fluid,  the  same  as  the  gall-bladder,  urinary 
bladder,  or  the  pelvis  of  the  kidney  and  ureter,  to  which  the 
structures  are  more  closely  approximated  ;  till  finally,  the  limit 
being  reached,  the  receptacle  contracts  for  compelling  it  out, 
driving  it  up  the  tube,  contraction  beginning  from  below  and 
continuing  on  up  the  canal  in  a  series  of  rhythmical  expan- 
sions and  contractions,  similar  to  what  occurs  in  the  oesopha- 
gus and  the  body  of  the  worms,  with  which  the  structures  are 
homologous,  the  same  law  applying  alike  to  all,  and  necessarily 
involving  similar  arrangements  in  the  structures  for  producing 
the  work.  In  the  present  case  we  have  the  great  abundance 
of  longitudinal  muscles  (Fig.  205,  c,  e)  for  effecting  shorten- 
ing in  the  tube,  thereby  diminishing  the  distance  and  reducing 
work  in  proportion,  while  the  circular  muscles  (d)  effect  the 
rhythmical  changes  in  pressure  for  compelling  movement,  the 
fluids  here,  as  elsewhere,  flowing  from  high  to  low  pressure  in 
conformity  with  organic  law.  In  other  words,  the  special 
adaptations  relate  to  work,  while  force  is  applied  in  the  measure 
of  the  requirements  (and  widely  removed  in  function  as  is  the 
local  vein  (g)  coursing  through  the  structures,  can  it  be  doubted, 
for  a  single  moment,  that  the  muscles  in  its  walls  also  relate 
to  work,  contracting  from  below  up  for  forcing  the  blood  to 
the  heart  and  lungs,  the  same  law  applying  to  both  ?). 

The  nerves  are  situated  in  the  tunica  adventitia,  and  form  a 
pretty  dense  plexus — the  plexus  spermaticus — in  a  portion 
distant  from  the  cremaster  internus,  and  are  derived  from  the 
spermatic  and  sympathetic.  From  the  plexus  spermaticus 
issue  several  smaller  nerve-trunks,  which  penetrate  the  mus- 
cular and  mucous  layers  of  the  vas  deferens,  where  they  are 
observed  to  have  medullated  fibres  (Klein).*  "In  the  upper 
portions  of  the  vas  deferens  small  ganglion  cells  are  scattered 
in  the  nerve-trunks  of  this  plexus,  and  also  in  those  trunks 
lying  more  externally  and  running  separately.  In  the  neigh- 
borhood of  the  ampullae,  however,  there  are  some  quite  feebly- 

*  Strieker's  Manual  of  Histology.  Art.,  Male  and  Female  External  Genital 
Organs  ;  together  with  their  Glandular  Appendages,  p.  584.     By  E.  Klein. 


496 


DUALISM  IN  NERVOUS  FORCE. 


developed  ganglion  cells."  It  will  thus  be  seen  the  nervous 
supply  to  the  organ  for  producing  and  coordinating  the  move- 
ments spoken  of  is  abundant. 


Jlt'l/Jlt  Ej'aetiUUry 
duct 


Fig.  207.— Base  of  the  Bladder  with  the  Vasa  Deferentia  and  Vesicula?  Seminales.- 

Gray. 


§bv^Olv 


Fig.  208.— Transverse  Section  through  the  Wall  of  a  Seminal  Vesicle.— Klein,  a, 
Epithelium  ;  6,  mucous  layer  ;  c,  internal  ;  d,  middle,  and  e,  outer  muscular  coat ; 
/,  adventitial  tunic  ;  g,  ganglia.     From  a  child. 

After  the  fluid  has  reached  the  seminal  reservoir,  it  is  still 
within  the  embrace  of  muscular  and  nervous  force,  the  ar- 
rangements which  obtain  in  the  parts  being  fundamentally  the 
same  as  in  the  vas  deferens,  only  that  the  cavitary  space  is 


DUALISM    IN    NERVOUS    FORCE.  497 

larger  and  the  rugse  in  the  mucous  lining  more  extensive  (Fig. 
208,  a),  the  muscles  (c,  d,  e)  not  quite  so  numerous,  compara- 
tively. Thence  it  is  compelled  into  the  urethra  within  the 
grasp  of  the  ejaculatory  muscles  during  the  sexual  orgasm, 
whence  it  is  driven  out  of  the  body  with  great  energy  under 
the  action  of  the  special  nerves.  The  large  nervous  ganglia 
(g)  in  the  adventitia  (/)  very  probably  serve  for  coordinating 
the  vas  deferens  with  the  action  in  the  walls  of  the  reservoir, 
as  also  for  compelling  emission,  in  this  manner  holding  similar 
relations  to  the.  ganglia  of  the  renal  plexus  for  producing  and 
coordinating  the  actions  in  the  kidney,  pelvis  and  ureter  for 
compelling  movement  in  the  contents,  the  matter  being  one  of 
variety  simply.  For  present  purposes  it  is  needless  to  extend 
the  matter. 

Striation  and  Fibrillation.— There  can  be  very  little  doubt 
that  striation  and  fibrillation  relate  to  electrical  tension  and 
increase  of  power  in  the  muscles  for  producing  energetic  ac- 
tion. In  the  slow-moving  snail,  for  example,  there  are  no 
striated  muscle  fibres  ;  but  in  the  earthworm  and  leech,  which 
possess  considerable  energy,  the  striated  are  freely  inter- 
spersed with  the  non-striated  muscle  fibres.  In  the  tongue  of 
the  frog,  in  which  the  action  is  very  energetic,  the  muscles  are 
strongly  striated  (Fig.  209). 

In  the  slow-moving  tortoise,  it  might  be  urged,  the  muscles 
are  also  striated  !  Yery  true  ;  but  it  is  no  exception,  and  ad- 
mits of  easy  explanation. 

Thus  for  lifting  and  transporting  the  heavy  house-like  body 
it  calls  for  the  expenditure  of  considerable  force,  hence  the 
striated  muscles  for  operating  the  bony  levers,  cervical  and 
caudal  vertebrae  ;  while  the  slow  movements  in  the  body 
result  from  the  defective  and  embarrassed  respiration,  pre- 
venting that  degree  of  oxygenation  of  the  blood  which  is 
essential  for  rapid  locomotion,  the  whole  forming  a  con- 
nected movement  in  the  very  nature  of  things,  for  evolving 
the  force  which  is  expended  in  the  activities,  as  has  already 
been  fully  set  forth;  hence  this  circumstance.  But  in  the 
individual  members  action  may  be  made  very  energetic,  nota- 
bly, the  retraction  of  the  head  and  limbs  when  the  parts  are 
irritated,  the  animal  doing  this  quickly  enough.     The  fishes 


498 


STRIATION   AND   NERVOUS   FORCE. 


move  rapidly  and  possess  great  muscular  power ;  the  muscles 
are  striated. 

In  short,  striation  in  muscles  corresponds  with  the  degrees  of 
energy  and  force.  This  circumstance  has  striking  illustration 
in  the  hollow  viscera  of  the  warm-blooded  animals,  notably 
the  heart,  terminal  portions  of  the  rectum,  the  ejaculatory  mus- 
cles in  the  urethra,  and  the  paunch  of  the  ruminant,  in  which 
the  muscles  are  striated,  whereas  in  all  the  other  viscera  the 
muscles  are  of  the  non-striated  variety,  for  the  reason  that 
rapid  action  is  not  called  for.     Finally,  in  embryognesis, 


Fig.  209.— Isolated  Muscular  Fibre  with  Transverse  Striae  from  an  Oblique  Section  of  the 
Tougue  of  a  Frog  Colored  with  Chloride  of  Gold.  The  muscle  cells  are  distinctly 
shown,  and  three  are  visible,  each  containing  several  nuclei.  P.  61  (Oc,  3  ;  obj.  8). 
Klein. 

in  which  development  repeats  itself,  we  have  the  transitional 
stages  in  muscle  evolution  also  manifesting  itself,  the  fusiform 
cells  of  non-striated  muscle  fibres  inosculating  and  under- 
going progressive  striation  (Fig.  210).  Then,  again,  we 
have  experimental  evidence  showing  electrical  phenomena 
in  the  muscles  during  musculation,  notably:  1.  Every  cross- 
section  of  a  striated  muscle  represents  a  negative  pole  of  an 
electric  current,  and  every  longitudinal  section  a  positive  pole ; 
and  it  is  immaterial  whether  the  section  be  made  with  a  knife 
or  corrosive  substance  ;  while  muscles  which  are  rigid,  or  which 


STRIATIC^  AND   NERVOUS   FORCE. 


499 


have  been  killed  without  "rigor"  being  induced,  exhibit  no 
current.  Du  Bois-Reymond  (Hermann).  2.  That  during 
the  passing  of  a  wave  of  contraction  along  the  bundle  of  fibres, 
produced  by  stimulating  the  end,  that  the  different  spots  in 


Fig.  210. — Muscular  Fibres  from  a  Foetal  Sheep  in  Process  of  Striation. — Frey.  a,  &, 
Very  long  fusiform  cells,  two  or  three  nuclei  and  commencing  striations  ;  c,  d,  por- 
tion of  a  somewhat  more  mature  fibre,  with  numerous  nuclei  and  considerable 
diameter  ;  e,  /,  g,  fibres  still  further  developed,  with  nuclei  in  the  axis  ;  h,  nuclei 
beneath  the  envelope  ;  i,  a  fibre  breaking  up  into  thick  disks. 

succession  on  the  longitudinal  surface  become  negative  in 
reference  to  other  spots,  there  being  a  negative  wave,  as  it 
were,  which  travels  along  at  the  same  rate  as  the  wave  of  con- 


500  STRIATION    AX1)    NEK  VOL'S   FORCE. 

traction,  viz.,  aboat  three  metres  per  second  (Bernstein'*)f 
"At  each  point  the  negative  state,  which  first  increases  and 
then  decreases,  lasts  about  vhs  of  a  second  ;  it  is  entirely  gone 
by  the  end  of  the  'latent  period,'  which  lasts  T*ff  of  a  second. 
Every  point  in  a  fibre  must  therefore  first  of  all  undergo 
electrical  changes  before  contracting  (Helmholtz,  Holmgren) ; 
or,  in  other  words,  the  wave  of  muscular  contraction  is  imme- 
diately preceded  by  a  negative  wave.  .  .  .  The  state- 
ment that  this  negative  wave  diminishes  in  intensity  as  it 
travels  on  (Bernstein)  appears  only  to  apply  to  the  case  of 
muscle  which  is  dying  (Du  Bois-Reymond)."  Italics  are 
added.  It  will  thus  be  seen  that  dualism  in  muscles  and 
nervous  force  is  further  corroborated  by  the  electrical  phe- 
nomena occurring  in  the  muscles  during  musculation,  the 
difference  in  the  currents  answers  to  the  opposite  molecular 
changes  in  the  sarcous  elements  corresponding  with  expan- 
sion and  contraction ;  otherwise  are  inexplicable.  But  the 
most  striking  circumstance,  the  fact  that  the  negative  wzve, 
which  answers  to  expansion,  precedes  the  positive,  which 
answers  to  contraction,  in  the  muscles ;  the  importance  of 
which  it  would  be  difficult  to  overestimate,  since  it  underlies 
the  mechanics  for  effecting  circulation,  so  that  diastole  in  the 
heart,  vessels,  and  hollow  viscera  is  not  only  in  conformity 
with  the  law  underlying  the  movement  in  the  fluids,  but  is 
further  made  inevitable  from  the  very  nature  of  the  mechanics 
in  musculation  ;  and  the  two  being  correlated  forces  in  nature, 
are  readily  coordinated  in  the  animal  functions  by  means  of 
the  nervous  apparatus,  which,  as  it  were,  separates  and 
limits  the  two  forms  of  electrical  force,  pouring  one  or  the 
other  into  parts,  so  as  to  produce  rapid  expansion  or  con- 
traction in  the  organs,  as  the  case  may  be,  in  the  exigencies  in 
the  functions ;  which  would  explain  the  special  phenomena, 
while  nothing  else  would.  In  short,  the  nervous  system  con- 
trols the  electrical  currents  in  the  muscles  ;  at  the  same  time, 
they  may  be  induced  artificially,  to  wit : 

*  Bernstein,  Untersuchungeri  iiber  d.  Erregungsvorgang  im  Nerven  und 
Muskelsysteme.     Heidelberg.    1871. 

f  Human  Physiology,  pp.  291-292.  By  L.  Hermann,  Professor  of  Physiology 
in  the  University  of  Zurich. 


STEIATION"  AND   JSTERVOl'S   FORCE.  501 

*  '•'In  a  perfectly  uninjured,  unskinned  animal  the  mus- 
cles, which  are  in  a,  state  of  rest,  are  entirely  free  from  elec- 
trical currents  (Hermannf)  ;  the  currents  originate  during  the 
preparation  of  the  muscle,  in  consequence  of  injurious  influ- 
ences acting  upon  their  surfaces.  In  frogs,  for  instance,  among 
other  such  influences,  is  to  be  mentioned  the  action  of  traces  of 
the  caustic  secretion  of  the  skin.  The  more  these  injurious 
influences  are  avoided,  the  greater  the  freedom  of  the  muscle 
from  electrical  currents.  In  muscles  which  are  at  rest  there 
are,  therefore,  no  currents  except  those  which  are  brought 
about  by  the  negative  electric  tension  of  the  artificial  cross- 
section  in  reference  to  the  longitudinal  section"  (Hermann). 

Then,  again  :  %  In  the  electrical  stimulation  of  muscles,  the 
same  laws  hold  as  in  that  of  nerves.  Here,  also,  it  is  only 
variations  in  currents  that  produce  stimulation,  which,  as 
before,  proceeds,  on  closing,  from  the  cathode,  and,  on  open- 
ing, from  the  anode  (von  Bezold). 

As  changes  take  place  more  slowly  in  muscle  than  in  nerve 
(as  evidenced,  for  example,  by  the  different  degrees  of  rapidity 
with  which  they  transmit  impressions),  length  of  duration  of 
the  stimulating  current  is  more  necessary  in  the  former  than 
in  the  latter  for  the  production  of  stimulation.  Hence,  all 
induction-currents,  and  the  more  transitory  constant  currents, 
are  unable  to  stimulate  to  contraction  muscle  deprived  of  its 
nervous  connections  by  curare,  while  they  are  able  to  cause 
contractions  in  a  muscle  by  acting  upon  its  motor  nerves 
(Brucke).  This  fact  was  early  known  to  be  the  case  with 
muscles,  the  nervous  organs  of  which  were  rendered  inca- 
pable of  performing  their  functions  by  exhaustion,  local 
death,  pathological  paralysis,  etc.  (von  Bezold,  Fick,  Neu- 
mann) "  The  above  brief  excerpts  will  be  sufficient  to  show 
the  nature  of  the  mechanics  in  musculation  and  nervous 
force.  Last,  but  not  least,  we  have  to  mention  the  sugges- 
tive fact  in  the  "striae"  themselves  (Figs.  211,  212),  which 
differ  essentially  from  each  other,  and  placed  one  above  the 

*  Hermann,  Weitere  Untersuchungen  uber  den  Stofftvechsel  in  Muskel.     Ber- 
lin, Verlag  v.  A.  Hirschwald.     1867. 
f  Hermann's  Physiology,  pp.  285-286. 
X Ibid.,  pp.  339-340. 


502 


STHIATIOX   AND   NKKVOUS   FORCE. 


other  alternately,  sustain  striking  resemblance  to  what  obtains 
in  a  Voltaic  pile  (Fig.  212,  a,  b),  the  sarcolemma  inclosing 
them  ;  also,  that  during  musculation  the  muscle  juices  give 
an  acid  reaction,  which  corresponds  with  the  conditions  in  an 
electric  pile.     Furthermore,  the  force  which  is  manifested  in 


Fig.  211.— Fragments  of  Striped  Elementary  Fibres,  showing  a  cleavage  in  opposite 
directions  ;  magnified  300  diameters. — Todd  and  Bowman.  A,  longitudinal  cleav- 
age—the longitudinal  and  transverse  lines  are  both  seen  ;  c,  fibrillar  separated  from 
one  another  by  violence  at  the  broken  end  of  the  fibre,  and  marked  by  transverse 
lines  equal  in  width  to  those  on  the  fibre  ;  c',  e",  represent  two  appearances  com- 
monly presented  by  the  separated  single  fibrillae  (more  highly  magnified)  ;  at  e'  the 
borders  and  transverse  lines  are  all  perfectly  rectilinear,  and  the  included  spaces 
perfectly  rectangular  ;  at  e",  the  borders  are  scalloped,  the  spaces  bead-like — when 
most  distinct  and  definite,  the  fibrilla  presents  the  former  of  these  appearances.  B, 
transverse  cleavage — the  longitudinal  lines  are  scarcely  visible  ;  a,  incomplete  frac- 
ture following  the  opposite  surfaces  of  a  disk,  which  stretches  across  the  interval  and 
retains  the  two  surfaces  in  connection — the  edge  and  surface  of  this  disk  are  seen  to 
be  minutely  granular,  the  granules  corresponding  in  size  to  the  thickness  of  the  diskr 
and  to  the  distance  between  the  faint  longitudinal  lines  ;  6,  another  disk  nearly 
detached  ;  b\  detached  disk,  more  highly  magnified,  showing  the  sarcous  elements. 


Fig.  2 IS. —Piece  of  Dead  Muscular  Filament  from  the  Fly.—  Engelmann.     a,  a,  Trans- 
verse disks  ;  b,  accessory  disks. 

the  muscles  during  energetic  action,  sufficient  to  rend  them 
asunder,  is  not  explicable  by  any  other  theory  than  the  one  of 
polar  action,  which  should  increase  cohesive  power  with  afflux 
of  electrical  force  ;  while  the  progressive  increase  in  hardness 
is  also  not  explicable  by  any  other  theory.  From  what  has 
preceded,  it  follows  :  1.  That  striation  and  fibrillation  relate  to 
increase  of  energy  in  the  muscles.      2.  That  two  forms  of 


GENESIS     OF    NEKVOUS    FOKCE.  503 

nervous  force  exist,  which,  answer  to  the  two  poles  in  the 
electric  current,  or  positive  and  negative  ;  while  the  principle 
for  producing  motion  in  muscles  is  by  changing  polarity  in  the 
molecular  elements,  which  is  the  function  of  the  nervous  appa- 
ratus. 

The  regular  intervals  and  correspondence  between  the 
striations,  the  transverse  markings  in  all  the  fibres  occur- 
ring at  the  same  intervals  (Fig.  211),  should  facilitate 
the  passage  of  the  nervous  currents  and  effect  concert  of 
action.  The  mode  of  nerve-terminations  also  throws  some 
light  upon  it.  Thus,  the  wall  or  sheath  of  the  nerve-tubes 
(Fig.  213,  B,  1, 1)  blends  with  the  sarcolemma  (3,  3),  with  which 
it  is  continuous,  the  terminal  nerve-plates  (5,  5)  being  within 
the  sheath  of  the  muscle-plates,  in  immediate  contact  with 
them,  the  medullary  substance  of  the  nerve-tubes  (4)  ceasing 
abruptly  at  the  site  of  the  nerve-plate,  whence  electrical  force 
is  diffused  through  the  muscle-plates  by  means  of  the  granular 
substance  which  forms  the  principal  part  of  the  nerve-plate, 
and  which  may  be  regarded  as  distributors  of  electrical  force, 
which  is  not  by  sharp  points,  but  blunt  or  spherical  bodies,  so 
as  to  more  uniformly  diffuse  it  through  the  fibre,  taking  from 
the  medullary  substance  and  transmitting  it  through  the 
muscle-plates.  But  here  again  comes  up  an  important 
question ;  notably :  ' '  What  becomes  of  the  force  which 
is  generated  in  the  muscles,  seeing  that  this  is  the  princi- 
pal seat  of  oxidation  and  source  of  body-temperature  ?  And 
while  heat  rapidly  diffuses  itself  through  the  tissues,  and  is 
removed  by  the  passing  blood,  we  know  that  considerable  pro- 
portion must  undergo  metamorphosis  into  electrical  force,  or 
similar  to  what  obtains  in  the  external  mechanics.  The  impor- 
tance of  this  question  will  at  once  be  seen  when  contrasted 
with  the  amount  of  work  performed  by  the  muscles,  which 
must  have  its  equivalent  in  electrical  force  derived  from  some 
source,  and  paid  for  there  and  then. 

Take  a  medium-sized  man,  or  one  of  one  hundred  and  fifty 
pounds,  for  example,  during  locomotion.  He  rises  to  the  erect 
position  and  steps  out — first  one,  then  the  other  leg,  following 
each  other  in  rapid  succession  in  locomotion  ;  the  force  which 
does  this  emanates  in  the  brain,  but  having  started  the  me- 


504 


GENESIS    OF    NERVOUS    FOKCE. 


clianics  fairly  and  well,  the  tiling  runs  itself  from  the  spinal 
cord  by  what  is  known  as  "reflex  action,"  propagated  through 
the  spinal  medulla,  the  brain  itself  having  little  to  do  with  it. 
Nor  can  it  be  imagined  for  a  single  moment  even  that  the 
force  for  producing  all  this  work  is  generated  in  the  cord  itself, 
the  special  nervous  centres  involved  being  far  too  inadequate 
for  this,  admitting  even  that  the  principle  was  right,  which, 
for  obvious  reasous,  can  by  no  means  be  done.     On  the  con- 


Fig.  213.— Mode  of  Nerve  Terminations  in  Striated  Muscles.—  Rouget.  A,  primitive 
fasciculus  of  the  thyro-hyoid  muscle  of  the  human  subject,  and  its  nerve  tube  ;  1,  1, 
primitive  muscular  fasciculus  ;  2,  nerve-tube  ;  3,  medullary  substance  of  the  tube, 
which  is  seen  extending  to  the  terminal  plate,  where  it  disappears  ;  4,  terminal  plate 
situated  beneath  the  sarcolemma,  that  is  to  say,  between  it  and  the  elementary 
fibrillar ;  5,  5,  sarcolemma.  B,  primitive  fasciculus  of  the  intercostal  muscle  of  the 
lizard,  in  which  a  nerve-tube  terminates  ;  1,  1,  sheath  of  the  nerve-tube  ;  2,  nucleus 
of  the  sheath  ;  3,  3,  sarcolemma  becoming  continuous  with  the  sheath  ;  4,  medullary 
substance  of  the  nerve-tube  ceasing  abruptly  at  the  site  of  the  terminal  plate  ;  5,  5, 
terminal  plate  ;  6,  6,  nuclei  of  the  plate  ;  7,  7,  granular  substance  which  forms  the 
principal  element  of  the  terminal  plate,  and  which  is  continuous  with  the  axis- 
cylinder  ;  8,  8,  undulations  of  the  sarcolemma  reproducing  those  of  the  fibrillse  ; 
9,  9,  nuclei  of  the  sarcolemma. 

trary,  the  work  involved  in  transporting  the  body  is  paid  for 
by  means  of  oxidation  of  fresh  materials  brought  into  the 
muscles  by  means  of  respiration  and  circulation,  which  are  in 
correspondence  with  the  activities,  so  that  the  force  is  neces- 
sarily generated  in  the  muscles  themselves ;  but  since  the 
nervous  apparatus  produces  and  regulates  the  muscular  move- 
ments, it  follows  that  the  force  so  generated  would  have  to 
pass  into  the  nerves  to  the  spinal  cord,  thence  out  again  into 
the  muscles,  the  nerves  serving  to  conduct  it  to  and  fro  be- 
tween the  muscles  and  the  ganglia,  under  guidance  and  direc- 


GENESIS     OF    NEKVO'S    FORCE. 


505 


tion  of  the  ganglia,  which  determine  the  movements.  In  short, 
it  wonld  seem  to  ns  that  the  electric  force  generated  in  the  tis- 
sues passes  np  one  set  of  nerves,  notably  the  so-called  "sen- 
sory nerves,"  to  the  posterior  columns  of  the  cord  (Fig.  214,  13), 
thence  through  the  gray  matter  of  the  cornua,  and  out  again 
through  the  anterior  roots  or  "motor  nerves"  (11,  11, 11)  from 
the  anterior  columns  to  the  muscles,  the  nervous  ganglia  in 
the  cord  serving  to  separate  and  limit  the  currents,  so  as  to 

A. 


Fig.  214. — Transverse  Section  of  the  Spinal  Cord  at  the  Origin  of  the  Fifth  Pair  of 
Cervical  Nerves. — Stilling.  In  this  figure,  the  white  substance  of  the  cord  is  repre- 
sented in  black,  to  show  more  clearly  the  limits  of  the  gray  matter  ;  1,  1,  antero- 
lateral columns  ;  2,  2,  posterior  white  columns  ;  3,  anterior  median  fissure  ;  4,  pos- 
terior median  fissure  ;  5,  white  commissure  ;  6,  gray  commissure  ;  7,  central  canal  ; 
8,  9,  anterior  cornua  of  gray  matter  ;  10,  10,  group  of  large  multipolar  cells  ;  11, 
11,  11,  anterior  roots  of  the  spinal  nerves  ;  12,  posterior  cornua  of  gray  matter  ;  13, 
posterior  roots  of  the  spinal  nerves. 

specialize,  at  the  same  time  regulating,  the  energy  of  the 
movements  in  the  muscles,  and  so  as  to  produce  expansion 
or  contraction,  as  the  case  may  be,  in  the  exigencies  in  the 
functions  ;  otherwise  impossible.  We  know,  too,  that  the 
nervous  apparatus  regulates  respiration  and  circulation  for 
evolving  the  force  ;  that  it  regulates  temperature  by  means  of 
the  vaso-motor  centre,  compelling  the  blood  to  the  skin  sur- 
face for  cooling  it,  which  would  involve  control  of  the  whole 
mechanics,  and  by  implication  the  force  generated  in  the  mus- 
cles ;  otherwise  it  would  be  utterly  impossible  to  produce, 
regulate  and   coordinate  the  movements  in  the  muscles,  as 


506 


GENESIS    OF    NEUVOUS    FOKCE 


must  appear  obvious.  Hence,  we  must  conclude  that  the  force 
generated  in  the  muscles  passes  up  to  the  cord  through  one 
set  of  nerves  and  down  again  through  the  other  set.  This 
principle  in  the  mechanics  has  further  corroboration  in  what 
occurs  in  the  case  of  the  sympathetic  ganglia  (Fig.  215),  in 
which  there  is  similar  arrangement,  the  nerves  passing  into 
the  nerve  knot  through  one  of  the  converging  trunks,  to  pass 
out  again  at  another  portion  of  the  knot.  Nay,  further,  the 
primitive  fibrils  pass  into  the  separate  cells  and  out  again 


Fig.  215.— Sketch  of  a  Mammalian  Sympathetic  Ganglion.—  Frey.  a,  b,  c,  Nervous 
trunks ;  d,  multipolar  cells  ;  d,  some  of  the  latter  with  a  dividing  nerve-fibre  ;  e, 
unipolar,  and  /,  apolar  cells. 

from  the  opposite  side,  forming  multipolar  cells  (d),  some 
of  the  latter  {d*)  with  a  dividing  nerve  fibre.  Why  some 
cells  (e)  have  only  one  fibre  is  explicable  only  by  the  hypoth- 
esis of  a  special  form  of  nervous  force  ;  while  the  apolar 
cells  (/)  would  refer  to  separating  and  disuniting  the  fluids, 
which  are  transferred  to  the  muscles  through  the  adjacent 
cells  and  nerves.  This  would  explain  the  presence  of  the  gan- 
glion on  the  posterior  root  of  the  spinal  nerves  (Fig.  113,  A\ 
with  the  fibres  running  into  and  out  of  the  cells,  thereby  re- 
lieving the  ganglia  in  the  spinal  cord  to  that  extent,  and  all 
sensory  nerves  are  thus  intersected  by  a  ganglion  knot.  The 
more  numerous  the  fibres,  the  larger  the  ganglion ;  notably 
the  fifth  and  the  eighth  pairs,  the  latter  possessing  actually 


GENESIS    OF    NERVOUS    FOKCE.  507 

two  such  ganglia.     And  being   the  universal  rule,  it  cannot 
be  doubted  for  a  single  moment  that  they  have  important  func- 
tions :    in  short,  tha,fc  they  effect  some  change  upon  the  nerv- 
ous fluid  ;  otherwise  are  meaningless.     This  circumstance  has 
forcible  illustration  in  disease  of  the  posterior  columns  of  the 
cord,  notably  "progressive  locomotor  ataxia,"  in  which  this 
transference  of  electric  fluid  through  the  nervous  arc  in  the 
cord  is  arrested,  and  every  movement  is  made  directly  from 
the  brain,  and  necessitating  the  concentrated  attention  and 
volition  of  the  patient  in  order  to  effect  locomotion,  which  is 
extremely  difficult,  showing  conclusively  that  the  main  source 
of  force  for  effecting  it  is  in  the  muscles  themselves  and  the 
changes  effected  upon  the  fluid  in  the  cord  itself.     In  short, 
it  underlies  the  principle  in  "reflex   action,"  which  means 
specialization  in  force ;  not  transformation,  but  rather  elec- 
tive   action,    separating    the    two    forms    of    electric    xorce ; 
otherwise,   this    also  would    be  meaningless.     Finally,    that 
there  is  such  transference  of  force — electric   force — from  the 
muscles  and  all  the  organs  to  the  appellate  nervous  centres, 
would  seem  inevitable,  from  the  very  nature  of  things.    Other- 
wise, the   comprehensive  arrangements  for  generating  force 
would  not  be  effective,  and  the  maintenance  of    a  balance 
utterly  impossible,  since  every  movement  involves   expendi- 
ture  of   force  for  producing  it.      One  other  thing  in   this 
connection,  and  we  will  bring  the  matter  to  a  close  ;  namely, 
the  very  intimate  relation  which  the  terminal  nerve  trunks 
sustain    to    each  other  in   the   tissues  (Fig.   216),  which  is 
not  by  inosculation,  as  Sir  Charles  Bell  thought  (the  finer 
work  attained  by  means  of  the  microscope  since  then  mak- 
ing this  known),   but  the  fibres  overlapping  simply,   which 
has  the  effect  of  completely  separating  the  currents,  enabling 
the  outgoing  current  to  pass    up  the    sensory  nerves    and 
the  in-going  down  the  other  or   motor  nerves,    without  in- 
terference, passing  readily  in  and  out  of  the  open  end  of 
the  nerves  through  the  terminal  plates  for  collecting  it.     And 
it  must  not  be  supposed  for  an  instant  that  the  brain  gene- 
rates  the  nervous  force  expended  in  the  organism,  though 
undoubtedly  it  contributes  greatly  to  it ;  but  as  in  the  case  of 
the  spinal  centres,   its   principal   role  is  the  separator  and 


508 


GENESIS    OF    NEEVOIS    FOKCE. 


regulator  of  the  nervous  supply,  and  a>  a  great  reservoir,  giv- 
ing it  out  in  the  measure  of  the  requirements.  .But  if  con- 
stantly drawn  upon  for  producing  the  voluntary  actions,  as,  for 
example,  in  walking  over  uneven  ground,  where  every  move- 


Fig.  216.— Nerves  of  the  Face,  fifth  and  seventh  pairs.— Sir  Charles  Bell.  A,  facial 
nerve  :  B,  trunk  of  same,  dissected  off  and  pinned  out ;  C,  branch  of  third  division  of 
fifth  nerve  joining  the  plexus  of  the  facial  ;  D,  masseteric  branch  of  fifth  nerve  ;  E, 
bucco-labial  branch  of  fifth  ;  F,  branch  of  fifth  to  muscles  of  lower  jaw  ;  G,  infra- 
orbital nerve  ;  H,  mental  nerve  ;  /,  infra-trochlear  nerve. 

ment  is  voluntary,  or  made  directly  from  the  brain,  it  speedily 
induces  exhaustion,  though  the  amount  of  work  actually  per- 
formed falls  far  short  of  that  done  upon  even  ground,  where 
the  movements  are  simply  reflex,  the  brain  itself  but  slightly 


GENESIS    OF    NERVOUS    FORCE.  509 

involved.  In  fine,  the  generation  of  force  involves  the  evolu- 
tion of  heat ;  hence  we  must  conclude  that  the  main  source  of 
force  in  the  organism  must  be  in  the  muscles  ;  nor  is  it  reason- 
able that  the  furnace  should  be  in  the  brain,  since  nervous  sub- 
stance can  endure  less  heat  than  any  other  tissue,  a  few  degrees 
of  blood  temperature  above  the  normal  amount  promptly 
arresting  its  functions,  inducing  prostration,  insensibility  and 
death.  But  the  nerves  terminating  in  the  muscles  as  they  do, 
the  neurilemma  blending  with  the  sarcolemma,  the  nerve-plates 
in  contact  with  the  muscle-plates,  we  can  readily  understand 
how  force  may  be  rapidly  generated  without  heating  the  brain, 
which  is  far  removed.  The  nerve  cells  being  the  most  highly 
organized,  we  can  readily  understand  the  special  arrangements 
that  obtain  respecting  them  and  their  relations  to  the  rest 
of  the  organism.  The  matter  needs  looking  into.  But  we 
may  rest  assured  that  everything  is  as  it  should  be  for  con- 
serving the  central  nervous  system,  at  once  the  seat  of  govern- 
ment and  throne  of  the  cell-brood. 

The  question  which  obtrudes  itself,  Can  force  be  evolved 
by  oxidation  in  the  tissues  without  producing  corresponding 
heat  %  has  deepest  import,  but  by  reason  of  the  nature  of  the 
problem  is  difficult  of  solution.  In  nerve  cells  heat  metamor- 
phosis into  electrical  force  is  probably  instantaneous  and  more 
or  less  complete,  else  active  mental  processes  would  be  fraught 
with  peril,  speedily  bringing  life  to  an  end  from  destructive 
changes  it  would  necessarily  superinduce  ;  hence,  we  must  con- 
clude that  heat  metamorphosis  is  more  or  less  complete  and 
instantaneous.  But  in  muscle  cells,  etc  ,  it  is  different ;  here 
an  amount  of  heat  is  evolved,  which  cannot  be  regulated 
in  any  other  way  than  by  bringing  the  blood  rapidly  to  the 
body-surface  for  cooling,  and  promoting  radiation  and  con- 
duction by  means  of  diaphoresis  and  evaporation,  at  the  same 
time  reducing  respiration  arid  circulation  for  diminishing  the 
importations,  as  has  already  been  fully  set  forth ;  otherwise, 
all  these  arrangements  would  be  meaningless.  Moreover,  it  is 
susceptible  of  easy  demonstration  by  simply  inhibiting  the  due 
escape  of  heat  from  the  body  for  maintaining  a  balance,  which 
is  promptly  followed  by  fever  and  constitutional  disturbance, 
and,  if   not  relieved,  would  undoubtedly  terminate  fatally. 


510  GENESIS   OF   NERVOUS   FORCE. 

Indeed,  one  may  be  speedily  aroused  from  deepest  sleep  by  in- 
creasing the  covering-  upon  Mm  simply,  the  sensations  distress- 
ful in  the  extreme,  the  imperious  necessity  and  dominant 
thought  to  cool  the  blood,  disrobing  with  all  impetuosity.  So, 
then,  we  must  conclude  that  oxidation  in  the  body-tissues 
necessarily  involves  evolution  of  heat,  and  that  the  arrange- 
ments spoken  of  for  regulating  temperature  are  the  only  means 
for  maintaining  a  balance  ;  the  only  question  being  whether 
there  is  exception  in  nerve  cells.  For  the  reasons  given, 
I  thirk  such  is  the  case.  While  life  is  an  "eddy"  in 
force,  the  central  nervous  system  is  the  heart  of  the  eddy 
where  force  culminates,  both  generated  in  itself,  and  rushing 
into  it  through  the  nerves  from  every'portion  of  the  body,  for 
evolving  the  movements  and  producing  the  psychical  phe- 
nomena, while  the  rapidity  of  THOUGHT  would  of  itself 
show  that  metamorphosis  is  instantaneous. 

THE  END. 


INDEX. 


PAGE. 

Abdomen,  a  ventricle 231 

Abdominal  sound 189 

Acineta,  action  in 55 

Actinosphaerum,  action  in 51 

Air,  atmospheric 1 

molecular  constitution  of 2,  5 

force  for  effecting  suspension  in 

space 7 

mode  of  producing  circulation  in, 

bo  as  to  form  the  currents   in    the 

earth 16,  17 

in  water 18 

bones 90 

in  intestines 187 

residual,  functions  of 123 

Air-bladders  in  fishes 373 

in  birds 80,  435 

Albumen,  rapid  absorption  of 80,  216 

Alveolar  action 122 

capillaries 123 

action  in 124 

Amoebae-movement,  import  of. .... .  46,  51 

mechanical  principle  for  effecting 

the  molecular  movements  in 447 

Animal  circulation,  principle  in 29,  32 

Arteries,  anatomy  of 148 

locomotion  of 151 

pyramids 153 

progressive  increase  of  the  muscles 

in 152 

development  of 436 

pulsations,  force  of 147 

Arterial  feeders  to  stomach 275 

small  intestines 276 

large  intestines 277 

Arterial  pressure  and  its  relation  to 

development 40,  41,  434 

physiological  problem  connected 

with  respiration 134 

Arterialization  of  the  blood  and  its  rela- 
tion to  development 41,  434 


PAGE. 

Automatism  in  the  vessels,  shown  in 

arterial  tracings 138 

Bile,  mode  of  circulation 228 

muscular  force  in 229 

automatism  in  the  gall-ducts 230 

Bladder,  urinary,  action  of 294 

gall,  action  of 231 

Blood,  circulation    of,    in    the    alve- 
oli  121,  136 

Brain,  oscillations  in 183 

Branchiae  in  arenicola 387 

terebella 389 

fishes 333 

Briinner's  glands 209 

Capillary  action,  automatism  in.  160,  168 

reflux  impossible 154 

Capillarity,  principle  in 19» 

Carbon,  atomic  weight 22- 

Carbonic  acid,  atomic  weight 10> 

normal  constituent  of  the  blood. . .  362 

function  of,  in  digestion 363 

in  the  nutritive  processes 369 

in  respiration 367 

not  a  poison 364 

a  food  for  the  tissues 366 

Cells,  columnar,  of  intestines 217 

Centre,  thermic 356 

rhythmic,  for  the  womb 335 

respiratory 32,  349 

vaso-motor 33,  349 

voluntary-motor 33,  349 

Chorion,  human  (eighth  week) 324 

Circulation,  primitive 52,  5$ 

in  amoebae 46,  55 

ditto,  gastrula 56 

ditto,  worms 383 

ground-plan  of 380 

planaria 381 

leech 383 

arenicola 387 

terrebella 389 


XX 


INDEX. 


PAGE.  , 

Circulation,  decapods 392 

lobster 394  | 

oyster 397 

fishes 399 

chick 344 

ditto,  tissue-interstices 176 

portal •  •  185 

hepatic 223 

embryonic 316,  436 

Coordination  of  internal  and  external 

parts  57,  60 

applied  to  respiration  and  the  ac- 
tion in  the  lungs 61,  65 

mode  of  effecting  it 67,  68 

stomach  and  walls  of  the  abdomen.  196 

Correlation  of  nervous  centres 32,  34 

Crocodile  connecting  link  with  the  birds  430 

Curves,  Traube's 137 

blood-pressure 145 

intrathoracic-pressure 134 

Defecation,  action  in 309 

Deglutition,  action  in 193 

Diaphragm,  office  of 86,    91 

adjustments  in 239 

action  of  the  pillars 240 

■ ditto,  vena  cava  lumen 243 

action  of,  in  defecation,  etc 309 

Dicrotism,  source  of 160 

multiple 172 

arrested 163 

Diffusion,  polar  action  in 20 

Digestion,  mechanical  action  in 204 

Dualism  in  muscles 456 

in  nervous  force 453,  503 

Ductus  venosus,  occasion  for 440 

Eel,  amphibious 407 

Electrical  force 7 

source  of 10 

r61e  of 7 

producing  circulation  in  the  at- 
mosphere       15 

producing  circulation  in  the  water.     16 

producing,     circulation     in     the 

plants 18, 19 

tension  in  muscles 453 

similarity  with  nervous  force 452 

generated  in  the  tissues 448 

Embryo,  an  acpiatic  animal 318 

human,  third  week 347 

Emesis,  action  in 315 

Erectile  tissue,  action  in 484 

Expansion,  principle  in 449 

demonstrated 452,  490 


PAGE. 

Expansion,  the  primary  movement.449, 500 

Fat,  rapid  absorption  of 218 

Fever,  physiology  of 253 

therapeusis  of 357 

Floor  to  the  viscera 84,  85,  300 

Floral  circulation,  principle  in 23,  27 

Foetal  circulation 442 

Force,  dualism  in  muscular  and  nerv- 
ous   444 

Frog,  circulation 416 

respiration,  principle  in 62 

Gall-ducts  230 

Gases,  functions  in  swim-bladders 373 

genesis  of 395 

ditto,  intestines 187,  362 

ditto,  stomach 203 

Gastric  capillaries 201 

action 204 

Gastrula,  action  in 56 

Glands,  peptic 198 

mucous 200 

Gullet,  action  of,  in  respiration 239 

Hardness  in  muscles  produced  by  nerv- 
ous force 461 

ameasure  of  work 452 

explanation  for 453 

Heart,  principle  in 126,  140 

relation  to  the  lungs 126 

ditto  tissues 153 

special  role  in 378 

beginning  of  cardial  development.  389 

great  relative   size  of  left  cham- 
bers, explanation  of 130 

not    the    force    in    the    circula- 
tion  174,  433 

tracings  deceptive 180 

tymph,  function  of 260 

embryonic 345 

first  evidence  in  terebella 389 

in  decapods 392 

ditto  lobsters 393 

oyster 397 

fishes 399 

development  of  left  auricle 412 

peculiarities    in     perennibranchi- 

ates 415 

in  the  frog 411 

in  chelonia 426 

interventricular  septum 428 

in  draco  volans 429 

in  crocodile 430 

in  birds 435 

Heat,  proportion  in  solar  beam 9 


INDEX. 


XXI 


PAGE. 

Heat,  metamorphosis  of,  into  electrical 
force  for  energizing  the  polar  forces.     10 

producing  circulation  in  air  and 

water  and  living  organisms 10,    22 

effect  upon  the  floral  circulation . .     24 

Intestines,    transverse    section   of,   in 

birds 187 

ditto,  mammals 187 

import  of  the  action  in,  or  peris- 
talsis     208 

physiological  anatomy  of 209 

capillaries  in 218 

glands  in 210 

Iris,  anatomy  of 477 

■ physiological  experiments  upon. . .  479 

Jaws,  mode   of  opening  and   closing 

them 470 

Kidneys,  physiological  anatomy  of 290 

■ nerves  of 293 

■ action  in 296 

Laughter,  pumping  action  in 225 

Law  in  the  animal  circulation 27 

Leech,  plan  of  the  circulation  in 385 

no  heart  in 384 

Lieberkuhn's  glands 210 

"  Lines  of  force  "  (Faraday) 11 

Liver,  anatomy  of 226 

circulation  in 227 

Lungs,  action  in 92,  100 

alveolar  collections,  mode  of  ex- 
pelling them 101,  102 

lung-action  demonstrated 109,  113 

circulation  of  air  in 92,  104 

ditto,  blood 121 

a  dual  circulation  in 35,  125 

mode  of  effecting  coordination 61 

nascent  condition 409 

in  chelonia 422 

molded  to  the  chest 114 

Lung-action  absolutely  necessary 99 

Lymphatics,  automatism  in 248 

physiological  anatomy 250 

vessels  of 252 

muscles 253 

valves 254 

commencement  of  lacfceals 255 

muscles  in 257,  258 

mechanical  action  in  the  walls  of 

intestine 256 

force  in  the  abdomen  for  increas- 
ing the  action 259 

hearts,  automatic  action  in 260 

Magnetism,  terrestrial 14,  15 


PAGE. 

Magnetism,  atmospheric 15,  16 

involved  in  polar  action 7 

molecular  movements  in  cells 447 

increased  by  electricity 453 

Mesentery 299 

action  in 298 

position  in  quadrupeds 85 

ditto,  man 300 

floor  of  support  to 84,  301 

Mucous  foldings  in  bronchia,  indica- 
tions of 101 

Muscles,  external  oblique,  action  in 302 

internal  oblique,  ditto 307,  308 

transversales,  ditto 305 

rectus  abdominis 305,  315 

levator  ani,  office  of 312 

of  the  tortoise 421 

cause  of  the  hardness  in,  during 

action 453 

stomach 206 

intestines 101 

levator  palpabrae 484 

orbicularis  oris 480 

palpabrarum 484 

penis 485 

Nerves,    recurrent   laryngeal,  unique 

distribution  accounted  for 118 

"inhibitor"    and    "accelerator," 

physiology  of 131,  283 

to  the  capillaries 133,  166 

ditto,  cell-brood ". .  178,  179 

to  the  viscera  in  the  abdomen 263 

correlation  of  nervous  force  in  . . .  264 

of  Meissner 265,  272 

■ ditto,  Auerbach Ibid. 

ditto,  columnar  epithelium. .  266,  270 

double  ganglionic  chain 278,  288 

roots  of  spinal  nerves  and  dorsal 

ganglia 279 

vaso-dilator  and  contractor 281 

ditto,  demonstrated 282 

pneumogastric,  action  in 284,  2S5 

— —  extensive  distribution  of 286,  287 

splanchnic 288,  289 

to  the  walls  of  the  abdomen 313 

ditto,  gravid  womb 333 

respiratory,  in  fishes 401,  406 

leech 59 

dualism  in 444,  503 

New-born,  changes  produced  by  birth .  338 
Nitrogen  gas,  functions  of  in  the  or- 
gans  370 

Nutrition,  principle  in 28 


xxn 


INDEX. 


PAGE. 

Nutrition,  rapidity  of,  in  plants 27 

difference  in  the  nutritive  proc- 
esses and  the  generation  of  force. .  27,  2S 

in  the  embryo 440 

below  the  diaphragm 439 

Odor,  f aecal,  source  of 3  <  1 

Osmose,  principle  in 20,  21 

Ovum,  human 348 

rabbit 323 

mole 348 

air-chamber  in 341 

physiology  of  the  circulation  in 

the  chick. 343 

Oxidation,  seat  of 359 

Oxygen  and  nutrition 439 

Oyster,  physiological  experiment  upon.  468 

Pancreas,  action  in 231 

Pelvic  viscera,  concentrating  force  in..  309 

Penis,  anatomy  of 485 

action  in 487 

Perineum,  floor  of 312 

Perennibranchiates,  circulation  in 415 

Physiological  problem  connected  with 
the  curves  of  blood-pressur*  and  intra- 
thoracic pressure  curves 134 

Placental  souffle,  analogue  of  respira- 
tion   319 

villi,  physiological  anatomy  of  —  326 

sinuses 327 

Pleuretic  adhesions 97 

Pneumatic  ducts 374 

Pneumonia,  mode  of  expelling  collec- 
tions  101 

Polar  force 6 

mode  of  energizing  polar  force 7 

action  in  the  atmosphere 11,  14 

action  in  the  water 18,  20 

action  in  diffusion 20 

action  in  osmose 21 

action  in  capillarity 21,  22 

action  in  floral  circulation 23,  25 

action  in  nutrition 27 

energy  of  in  flora 25,  26 

Portal  circulation,  physiological  experi- 
ments in 222 

dependence  upon  respiration 69 

Pressure,  its  relation  to  circulation. .  16,  17 

mechanics  for  effecting  changes  in 

pressure  in  the  atmosphere 15 

ditto  in  animal  organisms 29,  31 

arterial,  relation  to  animal  move- 
ment  40,  127,  156,  429 

fundamental  in  the  body 43,  45 


PAGE. 

Pressure  in  the  abdomen 189 

Pulsation,  first  visible  in  the  vessels 379 

Pulsus  venosus 128 

Respiration,       mechanical      principle 

in 35,36 

relation  to  venous  system 37,  38 

animal  movement 28,  497 

ditto  body-temperature 355 

two  respiratory  movements  per- 
forming at  the  same  time;  notably, 
one  in  the  lungs,  the  other  in  the  tis- 
sues      39 

in  different  stages  in  develop- 
ment     47 

frog 62,65 

bird 72 

mammalia 83 

in  the  tissues 160 

suspension  of,  in  deglutition 191 

effect  upon  the  portal  circula- 
tion  70,  220 

in  perennibranchiates 418 

ditto  chelonia 419 

axolotl 412 

fishes 405 

subaquatic 65,  424 

Respiratory  plane ...   303 

rocking  in  the  body 90 

Rete  mirabile,  office  of 377 

Rhythmic  centre  for  the  vessels 136 

lymphatics 248 

Sacral  promontory,  explanation  for 301 

Sacrum,  floor  to  pelvis 311 

Sighing,  physiology  of 224 

Sleep,  physiology  of 369 

Spleen,  anatomy  of 233 

action  in 235 

vein,  action  in 245 

Stoma :h  and  walls  of  the  abdomen  co- 
ordinated   196 

transverse   section    of,    showing 

mucous  fold 197 

physiological  anatomy  of ... .  198,  202 

gases,  secretion  of 203 

m3chanical  action 205 

muscles  of 206 

leech 386 

Sun,  source  of  electrical  supply 10 

producing  molecular  action  in  the 

atmosphere 8,  18 

ditto  water 19,  20 

ditto  plants 23,  26 

Temperature,  body 349 


IJSDEX. 


XXHL 


PAGE. 

Temperature,  genesis  of 350 

relation  to  musculation 351 

mode  of  maintaining  a  balance. . .  352 

Terebella,  plan  of  the  circulation  in, . .  390 

Thermic  centre 350 

Tidal  air 98 

Tissues,  respiration  in 160 

Tissue-circulation,  force  in 170 

Tongue  of  frog 134 

chameleon 486 

Trachea,  functions  in 104,  108 

physiological  experiments  on.  116,  117 

action  in  vocalization 120 

Tracings,  arterial 140 

artificial,  produced  by  apparatus 

of  Marey 141 

blood  pressure 145 

Umbilical  cord,  pulsations  in 147 

Uterine  sinuses,  pumping  action  in. .  326 

vessels 332 

centre  of  reflex  action 336 

contents,  mode  of  expelling 337 

Uterus,  physiological  anatomy  of 322 

impregnated 323 

Uterus  and  placenta,  section  of 325 

Vacuoles,  action  in 52 

Valves,  tricuspid,  insufficiency  of jl«» 

absence  of  in  portal  system 237 

lymph 254 

Vas  deferens 492 


PAGE. 

Vasiculae  seminales 496 

Vaso-dilator  nerves 277 

Vaso-contractor  nerves 278 

Veins,  physiological  anatomy  of 244 

Vense  cavae,  anatomy  of 244 

adjustments  in  inferior 243 

ditto,  superior 244 

mode  of  termination  at  the  heart.  436 

elongation  and  contraction  of 245 

Venous  ostia,  great  relative  size  of . . . .  129 

system,  action  in 183 

Vessels,  local  action  in 155 

maternal,  action  in 334 

changes  produced  by  birth 438 

automatism  in 139 

nervous  supply 157 

Villi,  anatomy  of 213 

Viscera  of  the  tortoise 426 

Vital  force,  nature  of 445 

Vocalization,  principle  in 118,  120 

Walls  of  the  abdomen,  section  of 315 

Water,  action  in 18 

composition  of 18 

in  perpetual  motion  from  changes 

in  polarity 19 

mechanics  in  evaporation  and  dif- 

rusion 20 

Waves,  respiratory  in  blood-pressure..  134 

arrested 173 

Work,  a  measure  of  force 452 


